Entomology 3rd edition - C.Gillott - Chapter 8 pps

8
The Hemipteroid Orders
1
. Intr
oduc
t
ion
T
he four orders (Psocoptera, Phthiraptera, Hemiptera, and Th
y
sanoptera) that constitut
e
t
he hemipteroid
g
roup are united b
y
the followin
g
features: specialized, usuall
y
sucto-
rial, mouthparts; small anal lobe in hind win
g
; win
g
venation reduced; cerci absent; fe
w
M
a
l

p
i
g
hi
an tu
b
u
l
es; an
d
ventra
l
nerve cor
d
w
i
t
hf
ew
di
screte gang
li
a. On t
h
ew
h
o
l
e, t
he

h
em
i
ptero
id
group
i
s more
h
omogeneous t
h
an t
h
e ort
h
optero
id
group, a
l
t
h
oug
h
two evo
l
u-
ti
onar
yli
nes

h
ave
d
eve
l
ope
d
,
l
ea
di
n
g
to t
h
e Psocoptera-P
h
t
hi
raptera, on t
h
e one
h
an
d
,an
d
t
he Hemiptera-Th
y

sanoptera, on the other
.
2
. Psocoptera
S
ynonyms:
C
orrodentia, Copeognatha
C
ommon names: barklice, booklice,
p
socids
S
mall or minute soft-bodied insects; mobile head with long filiform antennae and specialized
c
h
ew
i
n
g
mout
h
parts, compoun
d
e
y
es usua
lly
prom
i

nent
b
ut re
d
uce
di
n some spec
i
es; prot
h
orax
s
mall, win
g
s present or absent, le
g
s with two- or three-se
g
mented tarsi; external
g
enitalia of both
s
exes concealed
,
cerci absent.
This order, containin
g
about 3200 described species, has a worldwide, thou
g
h predom-

i
nant
l
y trop
i
ca
l
,
di
str
ib
ut
i
on. Some 290 spec
i
es occur
i
n Nort
h
Amer
i
ca, a
b
out 80
i
nBr
i
ta
i
n,

an
d 300 i
n Austra
li
a.
S
tructure
Psocoptera are stoc
ky
,so
f
t-
b
o
di
e
di
nsects w
h
ose
l
en
g
t
hi
s usua
lly l
ess t
h
an 10 mm

.
T
he lar
g
e, mobile head bears a swollen postcl
y
peus, lon
g
filiform antennae, and, usuall
y
,
prominent compound e
y
es, thou
g
h the latter are reduced in some win
g
less species. Three
oce
lli
are usua
ll
y present
i
nw
i
nge
df
orms
b

ut a
b
sent
i
n apterous spec
i
es. T
h
eY-s
h
ape
d
ep
i
cran
i
a
l
suture
i
s prom
i
nent. T
h
e mout
h
parts, t
h
oug
h

reta
i
n
i
ngac
h
ew
i
ng
f
unct
i
on, are
spec
i
a
li
ze
d
.T
h
e man
dibl
es are
di
ss
i
m
il
ar, t

h
ou
gh
eac
hh
as
b
ot
hg
r
i
n
di
n
g
an
d
cutt
i
n
g
e
dg
es.
I
n the maxillae the cardo and stipes are not alwa
y
s distinct. The
g
alea is a lar

g
e, flesh
y
lobe
,
1
99
200
CHAPTER
8
w
hereas the lacinia is a narrow, sclerotized rod (the pick), which ma
y
be used to scrape food
f
rom the substrate. The h
y
pophar
y
nx, which is able to take up water from the atmospher
e
(
Rudolph, 1982), has a characteristic structure. The lin
g
ua bears a pair of ventral sclerite
s
t
h
at are connecte
d

to t
h
eme
di
an s
i
top
h
ore sc
l
er
i
te
b
y
fi
ve
li
gaments. T
h
es
i
top
h
ore sc
l
er
i
t
e

i
ss
i
tuate
d
on t
h
e ventra
l
sur
f
ace o
f
t
h
e
b
ase o
f
t
h
ec
ib
ar
i
um. Oppos
i
te to
i
t, on t

h
e
d
orsa
l
sur
f
ace o
f
t
h
ec
ib
ar
i
um wa
ll
,
i
sa
k
no
blik
e process t
h
at
i
s
b
e

li
eve
d
to move a
g
a
i
nst t
he
sclerite in the manner of a mortar and pestle and facilitate the
g
rindin
g
up of food
.
In win
g
ed forms the small prothorax is lar
g
el
y
concealed b
y
the pterothorax. The win
g
s
are mem
b
ranous an
dh

ave a prom
i
nent
b
ut re
d
uce
d
venat
i
on. T
h
e anter
i
or pa
i
r
i
s
l
arger t
h
a
n
t
h
e
hi
n
d

pa
i
r. At rest t
h
ey are
h
e
ld
roo
flik
e over t
h
e
b
o
d
y. T
h
e
f
ore an
dhi
n
d
w
i
ngs are
c
oup
l

e
db
ot
hd
ur
i
ng
fli
g
h
tan
d
at rest. Vary
i
ng
d
egrees o
fb
rac
h
yptery occur, even w
i
t
hin
the same species, and apter
y
is common, especiall
y
in females. The le
g

s are usuall
y
slender
and similar; in some species the hind coxae carr
y
what is believed to be a stridulator
y
o
rgan. The abdomen is l0-segmented and terminates in a dorsal epiproct and a pair o
f
l
atera
l
paraprocts t
h
at may represent t
h
e 11t
h
segment. T
h
e externa
l
gen
i
ta
li
ao
f
ma

l
es
are wea
kl
y
d
eve
l
ope
d
,an
d
t
h
e
i
r
h
omo
l
og
i
es are uncerta
i
n. A sma
ll
ov
i
pos
i

tor
i
s usua
lly
present in females, thou
g
h it is much reduced or absent in some forms. Cerci are never
p
resen
t.
F
our Malpighian tubules originate at the posterior end of the midgut, which is long
an
d
convo
l
ute
d
.T
h
e nervous system
i
s
hi
g
hl
ymo
difi
e
d

an
d
compr
i
ses on
l
y
fi
ve gang
li
on
i
c
c
enters:
b
ra
i
n, su
b
esop
h
agea
l
gang
li
on, prot
h
orac
i

c gang
li
on, a compos
i
te pterot
h
orac
ic
g
an
g
lion, and a composite abdominal
g
an
g
lion. The tracheal s
y
stem usuall
y
opens to th
e
e
xterior b
y
means of two pairs of thoracic and ei
g
ht pairs of abdominal spiracles. Each
ov
ar
y

contains three to five pol
y
trophic ovarioles. The lateral oviducts are short and open
i
nto a
l
arger me
di
an
d
uct. A spermat
h
eca
i
s present. T
h
e testes are roun
di
s
h
or t
h
ree-
l
o
b
e
d
.
T

h
e vasa
d
e
f
erent
i
a
l
ea
di
nto
l
arge sem
i
na
l
ves
i
c
l
es t
h
at appear to pro
d
uce t
h
e mater
i
a

l
o
f
t
h
e spermatop
h
ore.
L
ife Histor
y
and Habit
s
M
ost Psocoptera are
f
oun
d
on vegetat
i
on or un
d
er
b
ar
k
,t
h
oug
h

some
li
ve among
l
ea
fli
tter, un
d
er stones, or
i
n caves. A
f
ew spec
i
es are assoc
i
ate
d
w
i
t
hh
umans an
d
may
be encountered in houses or buildin
g
s in which food materials are stored. Thou
g
h the

y
m
a
y
occur in vast numbers, the
y
are seldom of economic importance. The
y
are primaril
y
phytophagous, feeding on algae, lichens, fungi, pollen, and decaying fragments of highe
r
p
l
ants; occas
i
ona
ll
yt
h
ey eat
d
ea
d
an
i
ma
l
matter. Spec
i

es assoc
i
ate
d
w
i
t
hh
umans
li
ve on
c
erea
l
pro
d
ucts or,
i
nt
h
e case o
f
t
h
e common
b
oo
kli
ce, mo
ld

st
h
at
d
eve
l
op on o
ld b
oo
k
s
.
Man
y
species are
g
re
g
arious, with individuals of all a
g
es livin
g
to
g
ether, often beneath
a
silken web produced from secretions of the modified labial
g
lands. Most outdoor species
are fully winged, though brachyptery and aptery are common under certain environmenta

l
c
on
di
t
i
ons.
M
a
l
es are un
k
nown
i
n some spec
i
es, w
hil
e
i
not
h
ers
f
acu
l
tat
i
ve part
h

enogenes
i
sma
y
o
ccur. In
di
oec
i
ous spec
i
esama
l
et
y
p
i
ca
lly
courts a
f
ema
l
epr
i
or to mat
i
n
g
.E

gg
s,
b
etween
2
0 and 100 at a time, are laid sin
g
l
y
or in
g
roups, usuall
y
on ve
g
etation or under bark,
and the
y
ma
y
be covered with silk, particles of debris, or fecal material. The
y
have a thin
ch
or
i
on an
dl
ac
k

m
i
cropy
l
es an
d
aeropy
l
es. A
f
ew spec
i
es are v
i
v
i
parous. Larvae usua
lly
pass t
h
roug
h
s
i
x
i
nstars pr
i
or to metamorp
h

os
i
s,
b
ut t
hi
s
fi
gure
i
so
f
ten re
d
uce
di
npo
l
ymor-
p
hi
c spec
i
es. T
y
p
i
ca
lly
t

h
e apterous morp
h
s
h
ave one
f
ewer
i
nstar t
h
an t
h
e
f
u
lly
w
i
n
g
e
d
201
THE HEMIPTER
O
ID
ORDER
S
forms. Where climatic conditions permit, Psocoptera ma

y
have several
g
enerations per
y
ear
;
elsewhere, species are t
y
picall
y
univoltine and ma
y
enter diapause to overcome adverse
conditions
.
Phylogeny and
C
lass
ifi
cat
i
on
Modern Psocoptera are but the remnants of an order that had alread
y
under
g
one an
extensive evolution b
y

the end of the Permian period. Of the four hemipteroid orders, th
e
Psocoptera are genera
ll
y cons
id
ere
d
to
b
ec
l
osest to t
h
e ancestra
l
stoc
k
.T
h
e ear
li
est
f
oss
il
Psocoptera (
f
rom t
h

e Lower Perm
i
an o
f
Kansas [U.S.A.] an
d
Morav
i
a [Czec
h
Repu
bli
c])
diff
er
f
rom mo
d
ern spec
i
es w
i
t
h
re
g
ar
d
to w
i

n
g
venat
i
on an
d
mout
h
part c
h
aracters an
d
ar
e
placed in a distinct suborder Permopsocina. However, man
y
of the numerous Oli
g
ocene
fossils, and even some Cretaceous species, can be assi
g
ned to extant families (some eve
n
t
o extant genera) indicating that the order has undergone relatively little change since the
M
esozo
i
c. L
i

v
i
ng Psocoptera are
di
v
i
s
ibl
e
i
nto t
h
ree we
ll d
e
fi
ne
d
su
b
or
d
ers: Trog
i
omorp
h
a
,
T
roctomorp

h
a, an
d
Psocomorp
h
a (Eupsoc
id
a). T
h
e Trog
i
omorp
h
a conta
i
ns t
h
e most pr
i
m
-
itive and the Psocomorpha the most advanced Psocoptera. The Psocomorpha has been the
d
ominant suborder since the Late Cretaceous. Unfortunatel
y
, insufficient s
y
stematic work
h
as been done to

p
ermit firm conclusions to be reached with res
p
ect to the relationshi
p
so
f
t
he many families. This is especially true for the Psocomorpha, to which at least 7
5
%ofthe
recent species belong. The classification of Badonnel (19
5
1), which is used here, continues
t
o be accepted as the one that most accuratel
y
reflects ph
y
lo
g
enetic relationships within th
e
order
(
Smithers, 1991; Mockford, 1993
)
.
S
uborder Trogiomorph

a
D
i
st
i
n
g
u
i
s
hi
n
g
c
h
aracters o
f
Tro
gi
omorp
h
a
i
nc
l
u
d
e antennae w
i
t

h
more t
h
an 20 se
g
-
ments, never secondaril
y
annulated; tarsi three-se
g
mented; labial palps two-se
g
mented
;
pterosti
g
ma not thickened, or absent; and paraprocts with stron
g
posterior spine.
I
nt
hi
ssu
b
or
d
er t
h
ema
j

or
f
am
ili
es are t
h
e LEPIDOPSOCIDAE, TROGIIDAE, PSO
-
QUILLIDAE, and PSYLLIPSOCIDAE. Lepidopsocidae (16
5
species) form a primarily
t
rop
i
ca
lg
roup o
fb
ar
k
-an
dl
ea
fli
tter-
i
n
h
a
bi

t
i
n
gf
orms reco
g
n
i
ze
dby
t
h
e
i
r somew
h
at mot
h
-
like appearance produced b
y
the scales on their bod
y
and win
g
s. Thou
g
h a small famil
y
(about 20 species), the Tro

g
iidae is a cosmopolitan
g
roup that includes several species
found in buildings. Examples ar
e
Tr
o
g
ium pulsatoriu
m
(
Figure 8.1A), a common book
-
l
ouse t
h
at
f
ee
d
s on paper, vegeta
bl
e matter, an
d
cerea
l
pro
d
ucts, an

d
L
epinotus inqui
l
inus
th
at
i
s
f
oun
d
espec
i
a
ll
y
i
n granar
i
es an
d
ware
h
ouses. Psoqu
illid
ae an
d
Psy
lli

psoc
id
ae are
b
oth small (about 20 species in each), widel
y
distributed families. Psoquillids are found o
n
b
ark, in bird nests, and litter. Ps
y
llipsocids occur in caves and termite nests. Both families
include some s
p
ecies found indoors.
S
uborder Troctomorph
a
Features of Troctomorpha are 12- to 17-se
g
mented antennae, with some fla
g
ellar se
g
-
ments secondaril
y
annulated; 2- or 3-se
g
mented tarsi; 2-se

g
mented labial palps; pterosti
g
ma
not t
hi
c
k
ene
d
;an
d
paraprocts w
i
t
h
out a strong poster
i
or sp
i
ne
.
T
h
e
l
argest
f
am
ili

es
i
nt
hi
ssu
b
or
d
er are t
h
e LIPOSCELIDAE (140 spec
i
es)
,
AMPHIENTOMIDAE (40 species), and PACHYTROCTIDAE (
6
0 species). Th
e
202
CHAPTER
8
F
IGURE 8.1
.
P
socoptera.
(
A
)
Tro

g
ium pu
l
satoriu
m
(
Trog
iid
ae) (
di
sta
l
antenna
l
segments om
i
tte
d
); (B)
L
i-
posce
l
is
sp
.(L
ip
osce
lid
ae); an

d
(C
)
E
ctopsocus ca
l
ifornicus
(
Ecto
p
socidae). [A, from P P. Grass´e (ed.), 19
5
1
,
Tr
ait
r
r
ede
´
Z
oolo
g
ie,
V
ol. X. By permission of Masson, Paris. B, C, from L. A. Swan and C. S. Papp, 1972,
V
V
Th
e

C
ommon Insects of Nort
h
America.
C
opyr
i
g
h
t 1972
b
yL.A.Swanan
d
C. S. Papp. Repr
i
nte
db
y perm
i
ss
i
on o
f
Har
p
er&Row,Pu
bli
s
h
ers, Inc.]

L
i
posce
lid
ae
i
s a cosmopo
li
tan group w
h
ose mem
b
ers are recogn
i
ze
db
yt
h
e
i
r great
l
y en-
l
ar
g
ed hind femora. The famil
y
includes a number of common booklice (
L

i
p
osceli
s
s
pp.
)
(
Fi
g
ure 8.1B) found in houses, warehouses, and ship holds. Outdoor species t
y
picall
y
occu
r
i
n litter and under bark. However, occasionally they have been taken in the nests of ver-
te
b
rates,
i
nt
h
e
f
ur o
f
mamma
l

s, an
d
on
bi
r
d
s’
f
eat
h
ers, t
h
ese assoc
i
at
i
ons poss
ibl
ya
idi
ng
di
spersa
l
o
f
t
h
e psocopterans. Amp
hi

entom
id
s are pre
d
om
i
nant
l
y
f
oun
di
n trop
i
ca
l
reg
i
ons
o
f the Old World. The
y
occur under bark and in litter. Pach
y
troctidae are also mainl
y
trop
-
i
cal, occurrin

g
in both the Old and the New World. T
y
picall
y
, the
y
are found under bark o
r
i
n litter, occasionall
y
in buildin
g
s.
Suborder Psocomor
p
ha
F
eatures o
f
Psocomorp
h
a
i
nc
l
u
d
e antennae a

l
most a
l
wa
y
s 13-se
g
mente
d
,w
i
t
hfl
a
g
e
ll
ar
se
g
ments not secondaril
y
annulated; tarsi 2- or 3-se
g
mented; labial palps unse
g
mented;
pterosti
g
ma not thickened; and paraprocts with stron

g
posterior spine.
Ju
d
g
i
ng
b
yt
h
e
i
r common
f
eatures, t
hi
s very
l
arge su
b
or
d
er, conta
i
n
i
ng more t
h
an 20
f

am
ili
es,
i
s pro
b
a
bl
y
d
er
i
ve
df
rom or
h
a
d
a common ancestry w
i
t
h
t
h
e Troctomorp
h
a. Mos
t
m
em

b
ers o
f
t
h
esu
b
or
d
er are
f
oun
d
out
d
oors, on
g
row
i
n
g
ve
g
etat
i
on,
i
n
l
ea

fli
tter, or on
b
ar
k
.
The AMPHIPSOCIDAE (140 species) is a cosmopolitan famil
y
that includes some of the
l
ar
g
est Psocoptera. The
y
are
g
enerall
y
found on broad-leaved folia
g
e. ARCHIPSOCIDAE
(
60 species) form a tropical family, largely from South America and Africa, some species of
whi
c
hli
ve
i
n mass
i

ve aggregat
i
ons un
d
er s
h
eets o
f
we
bbi
ng t
h
at may cover an ent
i
re tree
.
T
h
e CAECILIIDAE
i
saver
yl
ar
g
e
f
am
ily
(370 spec
i

es most
ly i
nt
h
e
g
enu
s
C
aeci
l
iu
s
)
o
f
w
orldwide distribution. Most species are folia
g
e dwellers. ECTOPSOCIDAE (120 species)
,
w
hich form a cosmopolitan
g
roup, are t
y
picall
y
found in dr
y

folia
g
e and leaf litter thou
gh
af
ew
ff
sp
ecies are found elsewhere; for exam
p
le,
E
ctopsocus cali
f
ornicu
s
(
Figure 8.1C) is
a
co
mm
o
n
booklouse a
n
d
E. pumi
l
is
i

s a cosmopo
li
tan spec
i
es
f
oun
di
n granar
i
es an
d
ware-
h
ouses. PERIPSOCIDAE (120 spec
i
es) are
b
ar
kd
we
ll
ers w
i
t
h
a cosmopo
li
tan
di

str
ib
ut
i
on
.
Members of the ELIPSOCIDAE and EPIPSOCIDAE (each with about 100 species) are
203
THE HEMIPTER
O
ID
ORDER
S
b
ark and litter inhabitants, with worldwide (especiall
y
Southern Hemisphere) and primar
-
il
y
tropical distributions, respectivel
y
. The cosmopolitan LACHESILLIDAE (250 species
)
are primaril
y
found on dr
y
folia
g

e or in leaf litter with a few widespread species (e.
g
.,
Lac
h
esi
ll
ape
d
icu
l
ari
a
)
occurr
i
ng
i
n granar
i
es,
b
arns, an
d
ware
h
ouses. MYOPSOCIDAE
(12
5
species) are generally large, tropical Psocoptera found on bark. PHILOTARSIDAE

(1
5
0 species) live on bark or low ve
g
etation. The famil
y
is widel
y
distributed but especiall
y
common in the southwest Pacific re
g
ion. The PSEUDOCAECILIIDAE (120 species) is
a
mainl
y
tropical
g
roup of folia
g
e- and bark-dwellin
g
species. The PSOCIDAE, a cosmopoli-
t
an group of about
5
20 species, is the largest family in the order. Its members are darkly
co
l
ore

d
an
dli
ve on
b
ar
k
or, occas
i
ona
ll
y, on t
h
e groun
d
.
Literatur
e
A
d
eta
il
e
d
account o
f
t
h
e
bi

o
l
o
gy
o
f
t
h
e Psocoptera
i
s
gi
ven
by
New (1987) an
d
M
ockford (1993). Thornton (1985) discusses the zoo
g
eo
g
raph
y
and ecolo
gy
of the arborea
l
forms. The ph
y
lo

g
en
y
and classification of the order are considered b
y
Smithers (1972)
.
N
ew (1987) an
d
Sm
i
t
h
ers (1990) prov
id
e
k
eystot
h
ewor
ld f
am
ili
es an
d
genera, respect
i
ve
l

y
.
N
ew (1974)
h
as a
k
ey to t
h
eBr
i
t
i
s
h
spec
i
es, an
d
Sm
i
t
h
ers (1991) an
d
Moc
kf
or
d
(1993)

k
eystot
h
e Austra
li
an
f
am
ili
es an
d
Nort
h
Amer
i
can spec
i
es, respect
i
ve
l
y.
B
adonne1, A., 1951, Ordre des Psocopt`eres, in:
Tr
ait
r
r
ede
´

Z
oo
l
ogie (P P. Grass´e, e
d
.), Vo
l
. X, Masson, Par
i
s
.
M
oc
kf
or
d,
E. L.
,
1993
,
Nort
h
American Psocoptera (Insecta
)
,
San
dhill
Crane Press
,
Ga

i
nesv
ill
e
,
FL.
N
ew, T. R., 1974, Psocoptera, Han
d
b. I
d
ent. Br. In
s
ect
s
1
(7):1–102.
N
ew, T. R., 1987, B
i
o
l
ogy o
f
t
h
e Psocoptera, Or. Insects
21
:1
–

109.
R
u
d
o
l
p
h
, D., 1982, Occurrence, propert
i
es an
dbi
o
l
o
gi
ca
li
mp
li
cat
i
ons o
f
t
h
e act
i
ve upta
k

eo
f
water vapour
f
rom
t
he atmosphere in Psocoptera
,
J
.
I
nsect Ph
y
siol
.
2
8
:
111–121
.
S
m
i
t
h
ers, C. N., 1972, T
h
ec
l
ass

ifi
cat
i
on an
d
p
h
y
l
ogeny o
f
t
h
e Psocoptera
,
M
em. Aust. Mus
.
14
:1
–
349.
S
m
i
t
h
ers, C. N., 1990, Ke
y
stot

h
e
f
am
ili
es an
dg
enera o
f
Psocoptera
,
T
ec
h.
Rep. Aust. Mus
.
2
:
1–
8
2
.
S
mithers, C. N., 1991, Psocoptera, in
:
T
he Insects o
f
Australia
,

2nd ed., Vol. 1 (CSIRO, ed.), Melbourne Universit
y
P
ress, Car
l
ton, V
i
ctor
i
a
.
T
hornton, I. W. B., 198
5
, The
g
eo
g
raphical and ecolo
g
ical distribution of arboreal Psocoptera,
A
nnu. Re
v
. Entomo
l
.
3
0:175–196.
3

. Phth
i
ra
p
tera
S
ynonyms: Pseu
d
or
hy
nc
h
ota, Ma
ll
op
h
a
ga
C
ommon names:
li
ce, suc
ki
n
gli
ce (
i
n part),
(in part), Lipoptera (in part)
,

chewing lice or bird lice
Si
p
h
uncu
l
ata (
i
n part), (
i
n part)
Ano
p
lura (in
p
art)
Minute to small, apterous, dorsoventrally flattened ectoparasites of birds or mammals; head
prognat
h
ous or
h
ypognat
h
ous, compoun
d
eyes re
d
uce
d
or a

b
sent, oce
lli
a
b
sent, antennae 3
-
to
5
-se
g
mented, mouthparts of chewin
g
or piercin
g
-suckin
g
t
y
pe with palps of maxillae and
l
abium reduced or absent; prothorax free or fused with pterothorax, legs with unsegmented or
2
-se
g
mente
d
tars
i
, more or

l
ess mo
difi
e
df
or c
li
n
gi
n
g
to
h
a
i
ror
f
eat
h
ers, one, two, or no tarsa
l
claws; abdomen 7- to 10-segmented, cerci absent.
T
h
eor
d
er, w
hi
c
hh

as a wor
ld
w
id
e
di
str
ib
ut
i
on,
i
nc
l
u
d
es a
b
out 3100
d
escr
ib
e
d
spec
i
es
.
T
he suckin

g
lice (suborder Anoplura) are exclusivel
y
parasites of placental mammals,
w
hile the remainder are chewin
g
lice (“Mallopha
g
a”) comprisin
g
the suborders Ambl
y
cera
,
204
CHAPTER
8
Ischnocera, and Rh
y
nchophthirina. About 8
5
% of the species of chewin
g
lice are parasites o
f
birds. Man
y
Phthiraptera are important pests, in their role as disease vectors, of domesticated
animals and humans

.
S
tructur
e
T
he minute to small (0.35–10 mm lon
g
) bod
y
is dorsoventrall
y
flattened and shows
av
a
ried de
g
ree of sclerotization. In chewin
g
lice the head is relativel
y
lar
g
e and bear
s
ch
ew
i
ng mout
h
parts t

h
at s
h
ow certa
i
n resem
bl
ances to t
h
ose o
f
Psocoptera. In suc
ki
ng
li
c
e
t
h
ere
l
at
i
ve
l
y sma
ll h
ea
dh
as part

i
a
ll
y retracte
d
, suctor
i
a
l
mout
h
parts. T
h
e
l
a
b
rum
f
orms
a
s
h
ort evers
ibl
e pro
b
osc
i
s. T

h
ree st
yl
ets are conta
i
ne
d
w
i
t
hi
n a pouc
h
t
h
at runs ventra
lly
o
ff
the cibarium. The ventral st
y
let represents the modified labium, the middle st
y
let probabl
y
i
s an extension of the openin
g
from the salivar
y

duct, and the dorsal st
y
let is either th
e
m
odified maxillae or the hypopharynx. The mandibles disappear during embryogenesis
.
T
h
e
h
ea
db
ears t
h
ree- to
fi
ve-segmente
d
antennae t
h
at may
b
e
filif
orm (
i
n Anop
l
ura), cap-

i
tate an
di
n grooves (Am
bl
ycera), or mo
difi
e
df
or grasp
i
ng (Isc
h
nocera). Compoun
d
eyes
are reduced or absent and ocelli are never present. In Anoplura the thoracic se
g
ments ar
e
f
used, but in other lice the
p
rothorax is distinct from the
p
terothroax. The well-develo
p
ed
l
egs include a two-segmented or unsegmented tarsus and usually one or two tarsal claws.

E
i
g
h
t to ten v
i
s
ibl
ea
bd
om
i
na
l
segments occur. Ma
l
e gen
i
ta
li
a
i
nc
l
u
d
e a permanent
l
yev-
e

rte
d
en
d
op
h
a
ll
us. T
h
ere
i
s no true ov
i
pos
i
tor t
h
oug
hi
na
ll
suc
ki
ng an
d
some c
h
ew
i

n
g
l
ice the
g
onapoph
y
ses on se
g
ment 8 are used to attach e
gg
s to the host’s hair. Cerci ar
e
absent
.
In sucking lice the cibarium and pharynx form a strong sucking pump, but the crop
an
d
g
i
zzar
d
are poor
l
y
diff
erent
i
ate
d

.Inc
h
ew
i
ng
li
ce t
h
e crop
i
s
l
arge. In a
ll
P
h
t
hi
rapter
a
t
h
em
id
gut
i
s
l
arge an
dh

as two mesenter
i
c ceca. Four Ma
l
p
i
g
hi
an tu
b
u
l
es enter t
h
em
id
gu
t
poster
i
or
ly
.T
h
e nervous s
y
stem
i
s
highly

mo
difi
e
d
an
di
nc
l
u
d
es a compos
i
te metat
h
ora
-
c
oabdominal or thoracoabdominal
g
an
g
lion. The internal reproductive or
g
ans
g
enerall
y
resemble those of Psocoptera except that female Phthiraptera ma
y
have accessor

yg
land
s
an
dl
ac
k
as
p
ermat
h
eca
.
Lif
eH
i
story and Hab
i
t
s
Generall
y
, lice are considered to be hi
g
hl
y
host-specific bein
g
restricted to a sin
g

le o
r
af
ew
ff
closely related species of host, often occupying specific regions on the host’s body.
H
owever, Mars
h
a
ll
(1981) quest
i
one
d
w
h
et
h
er taxonom
i
c pro
bl
ems w
i
t
h
t
hi
s group may

b
eo
b
scur
i
ng t
h
e rea
l
s
i
tuat
i
on, c
i
t
i
ng t
h
e examp
l
eo
f
Menacant
h
us eur
y
sternu
s
,

w
hi
c
h,
o
n the basis of a recent revision, now counts 118 passerine species (in 20 families) and
5
w
oodpecker species amon
g
its hosts! B
y
contrast, a
g
iven bird or mammal ma
y
pla
y
host t
o
several species of lice, up to 15 being recorded on a South American tinamou. In contrast
to
fl
eas,
li
ce spen
d
t
h
e

i
r ent
i
re
lif
eont
h
e
h
ost. T
h
ey are
hi
g
hl
y sens
i
t
i
ve to c
h
anges
in
temperature an
dh
um
idi
ty an
d
surv

i
ve
f
or on
l
ya
f
ew
d
ays s
h
ou
ld
t
h
e
h
ost
di
e.
M
ost species of chewin
g
lice live amon
g
the feathers of birds where the
y
feed on
f
ra

g
ments of feathers and skin. A few species, for example, the chicken bod
y
louse
,
Me
n-
acanthus stramineu
s
, feed on blood in addition to e
p
idermal
p
roducts and are able to
p
ierc
e
t
h
es
ki
nor
d
eve
l
op
i
ng qu
ill
s. T

h
e mem
b
ers o
f
two sma
ll f
am
ili
es o
f
c
h
ew
i
ng
li
ce are para-
s
i
t
i
c on mamma
l
s, t
h
oug
h
t
h

e
i
r genera
lh
a
bi
ts are
lik
et
h
ose o
f
spec
i
es
f
oun
d
on
bi
r
d
s. Som
e
ch
ew
i
n
gli
ce,

lik
e Psocoptera,
h
ave t
h
ea
bili
t
y
to ta
k
eupmo
i
sture
f
rom t
h
e atmosp
h
ere v
i
a
205
THE HEMIPTER
O
ID
ORDER
S
t
heir h

y
pophar
y
nx, presumabl
y
an adaptation to prevent desiccation in the dr
y
air amon
g
feathers or fur of their host (Rudol
p
h, 1983).
The suckin
g
lice feed exclusivel
y
on the blood of the host, alwa
y
s a placental mammal.
I
t
h
as
b
een suggeste
d
t
h
at a poss
ibl

e reason
f
or t
h
e
hi
g
hh
ost spec
ifi
c
i
ty o
f
suc
ki
ng
li
ce
i
s
th
e
l
et
h
a
l
e
ff

ect an unsu
i
ta
bl
e
h
ost’s
bl
oo
d
m
i
g
h
t
h
aveont
h
e sym
bi
ot
i
c
b
acter
i
a present
in
certa
i

n
g
ut ce
ll
sor
i
nt
h
em
y
cetome, a structure c
l
ose
ly
assoc
i
ate
d
w
i
t
h
t
h
e
g
ut (see C
h
apte
r

16, Section 5.1.2
)
.
H
eav
y
infestations of lice ma
y
render a host more susceptible to disease and cause
econom
i
c
l
oss re
l
ate
d
to re
d
uct
i
on
i
n qua
li
ty.Ina
ddi
t
i
on, some suc

ki
ng
li
ce are
i
mportant
di
sease vectors (see Su
b
or
d
er Anop
l
ura).
I
n some Anop
l
ura, at
l
east, mat
i
ng occurs
f
requent
l
y, presuma
bl
y
b
ecause

f
ema
l
es
l
ac
k
a spermatheca. In man
y
species of chewin
g
lice, males are less common than females, and
in some species, are rare or unknown so that partheno
g
enesis occurs. E
gg
s are usuall
y
cemented to hairs or feathers by means of a secretion from the female’s accessory gland
.
Postem
b
ryon
i
c
d
eve
l
opment
i

s rap
id
,
j
uven
il
es pass
i
ng t
h
roug
h
t
h
ree mo
l
ts, an
d
a
d
u
l
ts
b
ecome sexua
ll
y mature w
i
t
hi

na
f
ew
d
ays o
f
t
h
e
fi
na
l
mo
l
t. Trans
f
er to new
h
osts
i
s
b
y
ph
y
sical contact and occurs durin
g
matin
g
, communal roostin

g
, and broodin
g
and feedin
g
t
he
y
oun
g
. Some bloodsuckin
g
Diptera ma
y
carr
y
lice from host to host
.
Phylogeny and Classification
I
tis
g
enerall
y
accepted that the Phthiraptera are derived from a free-livin
g
Psocopter
-
alike ancestor, based on a number of s
y

napomorphies (Chapter 2, Section 3.2). There i
s
virtuall
y
no fossil record: the Earl
y
Cretaceous chewin
g
louse
S
aurodectes vrsanskyi
may
h
ave paras
i
t
i
ze
d
pterosaur rept
il
es, an
d
a suc
ki
ng
l
ouse,
N
eo

h
aematopinus re
l
ictu
s
has been
f
oun
d
onaro
d
ent
,
C
ite
ll
us,
f
rom t
h
eP
l
e
i
stocene o
f
S
ib
er
i

a. However, t
hi
s
h
as not prevente
d
some aut
h
ors
f
rom specu
l
at
i
n
g
t
h
at t
h
eor
d
er ma
yb
equ
i
te anc
i
ent, poss
ibly

or
igi
nat
i
n
g
as
earl
y
as the Upper Carboniferous/Lower Permian. Kim and Ludwi
g
(1982) proposed thi
s
v
e
r
y
earl
y
ori
g
in for the order to be consistent with their h
y
pothesis that the
g
roup aros
e
f
rom an ancestor w
i

t
hi
nt
h
e psocopteran su
b
or
d
er Permopsoc
id
a, an
id
ea re
j
ecte
db
yLya
l
(198
5
). Rather, Lyal (198
5
) proposed, the lice may have evolved from liposcelidlike Pso
-
coptera t
h
at, as note
di
nt
h

e prev
i
ous sect
i
on,
h
ave occas
i
ona
ll
y
b
een
f
oun
di
nt
h
e nests an
d
on the bod
y
of birds and mammals. Initiall
y
, this association perhaps aided dispersal of the
insects, but in time the
y
ma
y
have be

g
un to feed on flakes of dead skin, bits of feathers, etc.
At this stage, the association between the ancestral louse and its host presumably woul
d
h
ave
b
een
f
acu
l
tat
i
ve
l
y paras
i
t
i
can
d
muc
hl
ess
h
ost-spec
ifi
ct
h
an

i
st
h
e case w
i
t
h
mo
d
er
n
P
h
t
hi
raptera. T
h
ev
i
rtua
l
a
b
sence o
ff
oss
il
s
h
as meant t

h
at proposa
l
sw
i
t
h
respect to t
h
eor
i
-
g
in and ph
y
lo
g
en
y
of the order have come lar
g
el
y
from examination of the host association
s
of lice, and the ph
y
lo
g
en

y
and zoo
g
eo
g
raph
y
of their mammalian and avian hosts. Suc
h
studies su
gg
est that ancestral lice did not arise until the Cretaceous, with either birds o
r
mamma
l
sas
h
osts. T
h
e ear
li
est
li
ce were c
h
ew
i
ng
f
orms, an

df
rom t
h
ese arose t
h
emo
d
ern
Am
bl
ycera on t
h
e one
h
an
d
,an
d
a
li
ne
l
ea
di
ng to t
h
e rema
i
n
i

ng groups on t
h
eot
h
er. T
h
e
l
atter
i
tse
lf
sp
li
t,
gi
v
i
n
g
r
i
se to t
h
e Isc
h
nocera, w
h
ose ancestor reta
i

ne
d
c
h
ew
i
n
g
mout
h
part
s
and had either a bird or a mammal as host, and the common ancestor of the Rh
y
ncoph
-
t
hirina and Anoplura which, like all modern members of these two
g
roups, presumabl
y
f
e
d
on mamma
l
s. W
hil
et
h

eR
h
yncop
h
t
hi
r
i
na reta
i
ne
d
c
h
ew
i
ng mout
h
parts, t
h
e Anop
l
ur
a
e
v
ol
ve
d
suctor

i
a
l
mout
h
parts
i
n con
j
unct
i
on w
i
t
h
t
h
e
i
r
bl
oo
d
-
f
ee
di
ng
h
a

bi
t. T
h
oug
h
t
hi
s
sc
h
eme ma
yb
e a reasona
bl
eexp
l
anat
i
on o
f
t
h
eevo
l
ut
i
on o
f
P
h

t
hi
raptera, Bar
k
er (1994
)
206
CHAPTER
8
F
IGURE 8.2
.
A
s
uggeste
d
p
h
y
l
ogeny o
f
t
h
eP
h
t
hi
raptera. T
h

ere
l
at
i
ons
hi
po
f
t
h
eTr
i
menopo
did
ae to ot
h
e
r
A
m
bly
ceran
f
am
ili
es
i
s uncerta
i
n.

has stressed that the coevolution of lice and their vertebrate hosts has not alwa
y
s occurred,
and that host-switching by lice is fairly common. Thus, a phylogeny based on lice character
s
rat
h
er t
h
an c
h
aracters o
f
t
h
e
h
ost
i
s pre
f
era
bl
e.
Some aut
h
ors p
l
ace t
h

ec
h
ew
i
ng
li
ce an
d
suc
ki
ng
li
ce
i
n separate or
d
ers, t
h
eMa
ll
op
h
ag
a
and Anoplura, respectivel
y
, the
g
reat differences in mouthpart structure and feedin
g

habit
s
bein
g
taken as sufficient
j
ustification for this separation. Generall
y
, however, all lice ar
e
i
ncluded in the order, Phthiraptera, divisible into four suborders, Amblycera, Ischnocera
,
R
h
yncop
h
t
hi
r
i
na, an
d
Anop
l
ura. T
h
us, t
h
eMa

ll
op
h
aga, w
hi
c
h
compr
i
ses t
h
e
fi
rst t
h
re
e
su
b
or
d
ers,
i
s a parap
h
y
l
et
i
c group

i
nt
hi
ssc
h
eme,
l
ea
di
ng some aut
h
or
i
t
i
es (e.g., Bar
k
e
r
et al.
,
2003) to ur
g
e that the term be abandoned. A su
gg
ested ph
y
lo
g
en

y
of the Phthirapter
a
i
s presented in Fi
g
ure 8.2
.
Su
b
or
d
er Am
bly
cer
a
M
embers of the suborder Ambl
y
cera have the followin
g
characteristics: capitate, four-
segmente
d
antennae,
l
y
i
ng
i

n grooves; man
dibl
es
h
or
i
zonta
l
, max
ill
ary pa
l
ps present; an
d
m
esot
h
orax an
d
metat
h
orax usua
ll
y separate
.
Generall
y
six families are reco
g
nized in this suborder, a

g
roup of about 8
5
0 species
that is considered to contain the more primitive lice. Of the six families, three are restricte
d
to avian hosts, two to marsu
p
ials, and one to
p
lacental mammals. The MENOPONIDAE
f
orm the largest family (650 species) and have a cosmopolitan distribution. Its members
i
n
f
est
bi
r
d
s, an
d
severa
l
spec
i
es are
i
mportant pests o
f

pou
l
try,
f
or examp
l
e, Menacant
h
u
s
s
tramineu
s
(t
h
ec
hi
c
k
en
b
o
d
y
l
ouse) an
d
M
enopon
g

a
ll
inae (t
h
es
h
a
f
t
l
ouse) (F
i
gure 8.3A).
The LAEMOBOTHRIIDAE are a small famil
y
containin
g
the sin
g
le
g
enu
s
Lae
m
obot
hr
io
n
,

207
THE HEMIPTER
O
ID
ORDER
S
F
IGURE 8.3.
P
hthiraptera. (A) The shaft louse, Menopon
g
allinae
(
Menoponidae); (B) the large turkey louse,
C
h
e
l
opistes me
l
eagri
d
is
(
Gon
i
o
did
ae
)

;
(
C
)
t
h
ere
d
catt
l
e
l
ouse
,
Dama
l
inia
b
ovi
s
(
Tr
i
c
h
o
d
ect
id
ae); (D) t

h
e
l
on
g
-
n
osed cattle louse
,
L
ino
g
nathus vitul
i
(Lino
g
nathidae); (E) the short-nosed cattle louse,
H
aematop
i
nus eur
y
sternus
(
Haematopinidae); (F) the human body louse
,
Pe
d
iculu
s

humanu
s
(
Pediculidae); and (G) the crab louse
,
P
hthiru
s
p
u
b
i
s
(P
h
t
hi
r
id
ae). [D, E,
f
rom L. A. Swan an
d
C. S. Pa
pp
1972,
Th
e Common Insects of Nort
h
America

.
C
op
y
r
igh
t
1972 b
y
L. A. Swan and C. S. Papp. Reprinted b
y
permission of Harper & Row, Publishers, Inc.]
th
e 14 spec
i
es o
f
w
hi
c
h
are paras
i
tes o
f
water
bi
r
d
san

dh
aw
k
s. T
h
e RICINIDAE, conta
i
n
i
ng
6
5
species in two
g
enera, are found on hummin
g
birds and several families of passerines. Be-
cause of the sister-
g
roup relationship of their hosts, the BOOPIIDAE (35 species), found o
n
Australian marsu
p
ials, and the TRIMENOPONIDAE (10 s
p
ecies), found on South Amer-
i
can marsup
i
a

l
san
dhi
str
i
comorp
h
ro
d
ents (porcup
i
nes, gu
i
nea p
i
gs, etc.), were
f
ormer
l
y
th
oug
h
tto
h
ave
h
a
d
a common ancestor. T

hi
sv
i
ew no
l
onger appears tena
bl
e; rat
h
er, t
he
B
oopiidae appear to be the sister
g
roup of the GYROPIDAE (
6
0 species). This famil
y
i
s
endemic to South and Central America, thou
g
h two species
,
G
yropus ovalis and
G
liricol
a
p

orcelli, which are found on
g
uinea pi
g
s, have been spread b
y
commerce to other parts of
th
ewor
ld
. Most spec
i
es o
f
Gyrop
id
ae
li
ve on
hi
str
i
comorp
h
ro
d
ents.
S
u
b

or
d
er Isc
h
nocera
Ch
aracter
i
st
i
cs o
f
Isc
h
nocera are expose
d
,t
h
ree- to
fi
ve-segmente
d filif
orm anten-
nae; man
dibl
es vert
i
ca
l
, max

ill
ar
y
pa
l
ps a
b
sent; an
d
mesot
h
orax an
d
metat
h
orax usua
lly
fused
.
208
CHAPTER
8
Included in this probabl
y
paraph
y
letic or even pol
y
ph
y

letic suborder of about 17
50
species are two lar
g
e families. The paraph
y
letic PHILOPTERIDAE, a cosmopolitan
g
rou
p
w
ith about 1460 species, is the lar
g
est famil
y
of the order. Its members are parasitic on
bi
r
d
san
di
nc
l
u
d
e a num
b
er o
f
pest spec

i
es
f
oun
d
on pou
l
try,
f
or examp
l
e, Cuc
l
uto
g
aste
r
h
etero
g
rap
h
us (t
h
ec
hi
c
k
en
h

ea
dl
ouse). T
h
e cosmopo
li
tan
f
am
il
y TRICHODECTIDAE
(
290 spec
i
es), w
hi
c
hi
s restr
i
cte
d
to p
l
acenta
l
mamma
l
s, conta
i

ns a num
b
er o
f
spec
i
es
f
ound on domesticated animals, for exam
p
le, Damalinia
(
Bovicola) bovis (cattle) (Fi
g
ur
e
8
.
3C)
,
D
.ov
is
(shee
p
), D. e
q
u
i
(

horses
),
F
elicola subrostratu
s
(
cats
)
, and Trichodectes canis
(d
ogs), w
hi
c
h
can serve as an
i
nterme
di
ate
h
ost
f
or t
h
e
d
og tapeworm,
D
ip
yl

i
d
ium caninu
m
.
T
h
e GONIODIDAE,
f
ormer
l
y
i
nc
l
u
d
e
di
nt
h
eP
hil
opter
id
ae, are
f
oun
d
on ga

llif
orm an
d
c
o
l
um
bif
orm
bi
r
d
s. T
h
e
f
am
il
y
i
nc
l
u
d
es some ma
j
or pou
l
try pests,
f

or examp
l
e,
Ch
e
l
o
p
iste
s
meleagridis
(
lar
g
e turke
y
louse) (Fi
g
ure 8.3B)
,
G
oniodes gigas (lar
g
e chicken louse) and
G
. dissimilis
(
brown chicken louse
).
Suborder Rhyncophthirina

M
embers of the suborder Rh
y
ncophthirina have the followin
g
characteristics: hea
d
prolon
g
ed into a rostrum, mandibles at apex of rostrum, and labium and maxillae vesti
g
ial
.
T
his suborder contains only two species, Haematomyzus elephantis
,
ap
arasite of bot
h
In
di
an an
d
A
f
r
i
can e
l
ep

h
ants, an
d
H.
h
op
k
insi
,
w
hi
c
hi
n
f
ests wart
h
ogs. It
h
as
b
een suggeste
d
t
h
at t
hi
s group may resem
bl
et

h
e ancestors o
f
t
h
e Anop
l
ura. Certa
i
n
l
y, t
h
e two groups s
h
ar
e
a number of primitive characters (L
y
al, 198
5
)
.
Su
b
or
d
er Ano
pl
ura

M
em
b
ers o
f
t
h
esu
b
or
d
er Anop
l
ura are recogn
i
ze
db
yt
h
e
i
rre
l
at
i
ve
l
y sma
ll h
ea

d
, sty
li-
f
orm mout
h
parts, an
dl
ac
k
o
f
man
dibl
es.
T
he approximatel
y
530 species of Anoplura were arran
g
ed b
y
Kim and Ludwi
g
(1978
)
i
n 15 families, of which 8 contain 4 or fewer species, The features of the seven lar
g
est fam-

ili
es, p
l
us t
h
ePe
di
cu
lid
ae an
d
P
h
t
hi
r
id
ae, w
hi
c
hh
ave spec
i
a
l
s
i
gn
ifi
cance

f
or
h
umans, are
summar
i
ze
db
e
l
ow , T
h
e ECHINOPHTHIRIIDAE (12 spec
i
es) are paras
i
tes o
f
P
i
nn
i
pe
dia
(
sea
l
s, sea
li
ons, an

d
wa
l
ruses
)
an
d
t
h
er
i
ver otte
r
(Lutra cana
d
ensis)
.
T
h
e
i
r
b
o
d
y
i
s covere
d
w

ith stron
g
spines or scales that retain a film of air over the bod
y
when submer
g
ed. It i
s
e
vident that man
y
of these lice must be ver
y
lon
g
-lived, as their hosts spend most of their
l
ife at sea, coming ashore to breed (when presumably transfer of parasites occurs) for onl
y
as
h
ort t
i
me eac
h
year. T
h
e cosmopo
li
tan LINOGNATHIDAE (70 spec

i
es) paras
i
t
i
ze
d
ogs
(
Lino
g
nat
h
us setosus
)
,h
y
raxes, an
d
rum
i
nants,
i
nc
l
u
di
ng s
h
eep

(
L. ovi
ll
u
s
a
n
d
L
.
p
e
d
a
l
is)
,
c
attle
(
L. vituli
)
(
Fi
g
ure 8.3D), and
g
oat
s
(L. steno

p
sis).
T
he HAEMATOPINIDAE
(
22
s
pecies) form a cosmopolitan
g
roup whose members parasitize un
g
ulates, includin
g
several
domesticated forms on which the
y
ma
y
become serious pests
.
Haematop
i
nus eurysternus
(
F
i
gure 8.3E) occurs on catt
l
e,
H

.
s
u
is
o
np
i
gs, an
d
H
.a
si
n
i
on
h
orses. T
h
eH
O
PL
O
PLE
U
RI-
DAE
(a
b
out 130 spec
i

es) are a
l
so a
l
arge group w
h
ose
h
osts are ma
i
n
l
yro
d
ents an
d
ra
bbi
ts
,
b
u
t
a
l
so
i
nc
l
u

d
emo
l
es an
d
s
h
rews. T
h
e
l
ar
g
est anop
l
uran
f
am
ily i
st
h
e POLYPLACIDAE
,
a cosmopolitan
g
roup whose 175 species ver
y
lar
g
el

y
parasitize rodents, thou
g
h some are
f
ound on primates (lemurs, lorises, and
g
ala
g
os), rabbits, moles, and shrews. The ENDER
-
LEINELLIDAE (50 species) are another rodent-infesting group, specifically parasitizin
g
S
c
i
ur
id
ae (squ
i
rre
lf
am
il
y). T
h
e
f
am
il

y
i
sw
id
esprea
d
t
h
oug
h
not represente
di
n Austra
li
a,
Mada
g
ascar, and southern South America. PEDICINIDAE (1
6
species) infest Old Worl
d
209
THE HEMIPTER
O
ID
ORDERS
monke
y
s. The PEDICULIDAE (2 species) live on primates. The
g

enus
P
e
d
iculu
s
i
s foun
d
on humans
(
a
s
P.
humanus
a
n
d
P.
c
a
pi
t
i
s
)
and other hominoids as well as New World mon
-
ke
ys

.
P.
c
ap
i
ta
s
(the head louse) and
P.
h
umanus
(
the bod
y
louse) (Fi
g
ure 8.3F) differ i
n
s
i
ze an
dh
a
bi
ts. T
h
e sma
ll
er
h

ea
dl
ouse attac
h
es
i
ts eggs to
h
a
i
r, w
h
ereas t
h
e
b
o
d
y
l
ouse
or “coot
i
e”
l
ays
i
ts eggs on c
l
ot

hi
ng to w
hi
c
hi
t usua
ll
y rema
i
ns attac
h
e
d
. Unt
il
recent
l
y
,
P
.
h
umanu
s
a
n
d
P
.
ca

p
itis
w
ere cons
id
ere
d
s
ibli
n
g
spec
i
es. However, a recent mo
l
ecu
l
ar
stud
y
(Le
o
et al.
,
2002) indicated that head and bod
y
lice are conspecific, despite their
d
ifferences. Certainl
y

, the
y
interbreed readil
y
in the laborator
y
. This discover
y
is si
g
nifi-
cant g
i
ven t
h
at t
h
etwo
f
orms are genera
ll
y
b
e
li
eve
d
to
b
e very

diff
erent
i
nt
h
e
i
rro
l
es a
s
di
sease vectors. T
h
us, t
h
e
b
o
d
y
l
ouse
i
san
i
mportant vector o
fdi
seases suc
h

as en
d
em
ic
t
yp
h
us an
d
trenc
hf
ever, cause
db
y
bl
oo
d
-
b
orne r
i
c
k
etts
i
as. Re
l
aps
i
ng

f
ever, cause
db
ya
spirochete, is also spread b
y
the louse. B
y
contrast, the head louse is not usuall
y
consid-
ered a disease vector
(
but see Robinso
n
et al.
, 2003
)
. Included in the PHTHIRIDAE ar
e
Phthirus gorillae, found on gorillas, and
P.
pubi
s
, the pubic or crab louse (Figure 8.3G)
f
oun
d
on
h

umans. Un
lik
et
h
e
b
o
d
y
l
ouse, t
h
e cra
bl
ouse
d
oes not appear to
b
ea
di
sease
vector
.
Literatur
e
I
n
f
ormat
i

on on t
h
e
bi
o
l
ogy o
f
P
h
t
hi
raptera
i
sg
i
ven
b
yAs
k
ew (1971), Mars
h
a
ll
(1981)
,
and Price and Graham (1997). Ferris (19
5
1) and Kim (198
5

) deal specifically with the
Anoplura (thou
g
h the former’s s
y
stematic treatment of the
g
roup is outdated), while Emer-
son and Price (1985) consider the Mallopha
g
a on mammals. Cla
y
(1970), L
y
al (1985)
,
B
arker (1994), Cruickshank
et al.
(
2001), and Barke
r
et al.
(2003) discuss the phylogeny o
f
th
eor
d
er. Arnett (2000)
i

nc
l
u
d
es
k
eys to Nort
h
Amer
i
can
f
am
ili
es o
f
P
h
t
hi
raptera, Ca
l
a
by
an
d
Murray (1991) to t
h
e Austra
li

an
f
am
ili
es o
f
P
h
t
hi
raptera, an
d
K
i
man
d
Lu
d
w
i
g (1978) to
w
or
ld f
am
ili
es o
f
Anop
l

ura. K
i
m
et al.
(198
6
) provide a ke
y
for North American Anoplura
.
A
rnett, R. H., Jr.,
2000,
A
merican Insects: A Han
db
oo
k
of t
h
e Insects of America Nort
h
of Mexico,2n
d
e
d
., CRC
P
ress
,

Boca Raton
,
FL
.
A
skew, R. R., 1971
,
P
ar
asitic Insects
r
r
, American Elsevier, New York
.
B
ar
k
er, S. C., 1994, P
h
y
l
ogeny an
d
c
l
ass
ifi
cat
i
on, or

i
g
i
ns, an
d
evo
l
ut
i
on o
fh
ost assoc
i
at
i
ons o
fli
ce
,
Int. J. Parasito
l.
24
:128
5
–12
9
1.
B
arker, S. C., Whiting, M., Johnson, K. P., and Murrell, A., 2003, Phylogeny of the lice (Insecta: Phthiraptera
)

i
n
f
erre
df
rom sma
ll
su
b
un
i
t rRNA, Zoo
l
. Scrip.
3
2:
40
7–
414
.
Ca
l
a
by
, J. H., an
d
Murra
y
, M. D., 1991, P
h

t
hi
raptera,
i
n: T
h
e Insects of Austra
l
ia
,
2n
d
e
d
., Vo
l
.I
(
CSIRO, e
d
.
)
,
M
elbourne University Press, Carlton, Victoria
.
C
l
ay, T., 1970, T
h

eAm
bl
ycera (P
h
t
hi
raptera: Insecta), Bu
ll
. Br. Mus. Nat. Hist.
25
:75–98.
Cru
i
c
k
s
h
an
k
, R. H., Jo
h
nson, K. P., Sm
i
t
h
, V. S., A
d
ams, R. J., C
l
a

y
ton, D. H., an
d
Pa
g
e, R. D. M., 2001,
P
hylogenetic analysis of partial sequences of elongation factor
1
α
i
dentifies major groups of lice (Insecta
:
Ph
t
hi
raptera)
,
M
o
l
.P
h
y
l
og. Evo
l.
19
:
202–215

.
E
merson, K. C., and Price, R. D., 198
5
, Evolution of Mallopha
g
a on mammals, in:
C
oevo
l
ution of Parasiti
c
A
rthro
p
ods and Mammal
s
(K. C. Kim, ed.), Wiley, New York.
F
erris, G. F., 1951, The sucking lice, Mem. Pac. Coast Entomo
l
. Soc.
1
:
320
pp
.
Kim, K. C., 198
5
, Evolution and host associations of Ano

p
lura, in: Coevo
l
ution of Parasitic Art
h
ropo
d
san
d
M
ammal
s
(
K. C. Kim, ed.), Wiley, New York
.
K
i
m, K. C., an
d
Lu
d
w
i
g, H. W., 1978, T
h
e
f
am
il
yc

l
ass
ifi
cat
i
on o
f
t
h
e Anop
l
ura,
S
yst. Entomo
l.
3
:249
–
284.
K
i
m, K. C., an
d
Lu
d
w
ig
, H. W., 1982, Para
ll
e

l
evo
l
ut
i
on, c
l
a
di
st
i
cs, an
d
c
l
ass
ifi
cat
i
on o
f
paras
i
t
i
c Psoco
d
ea
,
Ann.

Entomol.
S
oc. Am
.
75
:
53
7–
5
4
8
.
Kim, K. C., Pratt, H. D., and Stojanovich, C. J., 198
6
,
Th
e Suc
k
ing Lice of Nort
h
America: An I
ll
ustrate
d
Manua
l
f
or I
d
enti

fi
catio
n
, Penns
yl
van
i
a State Un
i
vers
i
t
y
Press, Un
i
vers
i
t
y
Par
k.
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8
Leo, N. P., Campbell, N. J. H., Yan
g
, X., Mumcuo
g
lu, K., and Barker, S. C., 2002, Evidence from mitochondrial
DNA that head lice and body lice of humans (Phthiraptera: Pediculidae) are conspecific

,
J. Med. Entomol.
39
:66
2–
666.
L
yal, C. H. C., 1985, Phylogeny and classification of the Psocodea, with particular reference to the lice (Psocodea:
L
L
Phthiraptera)
,
Syst. Entomol.
10
:145–165
.
Mars
h
a
ll,
A. G.
,
1981
,
Th
e Eco
l
ogy of Ectoparasitic Insect
s
,

Aca
d
em
i
c Press
,
New Yor
k.
P
rice
,
M. A.
,
and Graham
,
O. H.
,
1997
,
C
hewing and Sucking Lice as Parasites o
f
Mammals and Birds
,
U.S.D.A.
,
Tec
h.
Bu
ll.

1
84
9
:
1
–
309
.
R
o
bi
nson
,
D.
,
Leo
,
N.
,
Proc
i
v
,
P.
,
an
d
Bar
k
er

,
S. C.
,
2003
,
Potent
i
a
l
ro
l
eo
fh
ea
dli
ce
,
P
e
d
icu
l
us
h
umanus capiti
s
,
a
s
v

ectors of Rickettsia
p
rowazeki
i
,
Par
asitol. Res.
rr
90
:
20
9
–211
.
R
u
d
o
l
p
h
, D., 1983, T
h
e water-vapour upta
k
e system o
f
t
h
eP

h
t
hi
raptera, J
.
Insect P
h
ysio
l
.
29
:15–25.
4
. Hem
ip
ter
a
Synonym:
Rhy
nc
h
ot
a
C
ommon name: true
b
u
gs
M
i

nute to
l
ar
g
e
i
nsects,
h
ea
d
op
i
st
h
o
g
nat
h
ous (
h
omopterans) or pro
g
nat
h
ous (
h
eteropterans),
c
ompound eyes usually well developed but rarely absent, two or three ocelli usually present,
a

ntennae w
i
t
hf
ew se
g
ments, mout
h
parts suctor
i
a
l
w
i
t
h
man
dibl
es an
d
max
ill
ae
i
n
f
orm o
f
st
yl

et
s
e
nclosed within a labial sheath; two pairs of win
g
s usuall
y
present with fore win
g
s of harder
c
onsistency than hind pair; abdomen with 9–11 segments, external genitalia varied in both sexes,
c
erc
i
a
b
sent
.
Thi
s group
i
st
h
e
l
argest an
d
most
h

eterogeneous or
d
er o
f
exopterygotes, conta
i
n
i
ng
some 80,000
d
escr
ib
e
d
spec
i
es
f
rom a
ll
re
gi
ons o
f
t
h
ewor
ld
,A

b
out 11,000 spec
i
es
h
ave
been described from North America
,
5600 from Australia
,
and 1600 from Britain. The
o
rder traditionall
y
has been divided into the Homoptera (Sternorrh
y
nch
a
+
A
uchenorrh
y
n-
ch
a) an
d
Hetero
p
tera (an
d

some aut
h
ors
h
ave ra
i
se
d
t
h
ese to t
h
e ran
k
o
f
or
d
er). How-
ev
e
r, t
h
ere
i
s a grow
i
ng
b
o

d
yo
f
ev
id
ence,
b
ot
h
morp
h
o
l
og
i
ca
l
an
d
mo
l
ecu
l
ar, to
i
n
di
cat
e
t

h
at t
h
e Homoptera
i
s a parap
hyl
et
i
c
g
roup, an
d
t
h
at t
h
e Auc
h
enorr
hy
nc
h
a
i
st
h
es
i
ste

r
g
roup of the Heteroptera (see Ph
y
lo
g
en
y
and Classification). Thus, in the description be
-
l
o
w
the ter
m
homoptera
n
is used to describe a level of or
g
anization, not a taxonomi
c
ran
k
.
S
tructur
e
Hem
i
ptera range

i
ns
i
ze
f
rom a
b
out1mmto11cm.T
h
ey are
f
requent
l
y procrypt
i
ca
ll
y
c
olored in shades of
g
reen or brown, or aposematicall
y
colored in strikin
g
patterns, ofte
n
i
n red and white. Yet others mimic ants and other insects with which the
y

live, or resem-
ble plant parts. Various forms of polymorphism (see Chapter 21, Section 7) are commo
n
t
h
roug
h
out t
h
eor
d
er. T
h
e
h
ea
di
se
i
t
h
er op
i
st
h
ognat
h
ous an
dl
ac

ki
ngagu
l
a
i
n
h
omopteran
s
o
r prognat
h
ous an
d
w
i
t
h
asc
l
erot
i
ze
d
gu
l
a
i
n most Heteroptera. T
h

e antennae compr
i
se on
l
y
f
our or five se
g
ments in most Hemiptera. The compound e
y
es are t
y
picall
y
well develope
d
and of varied shape and size. Two or three ocelli are usuall
y
present. The unif
y
in
g
feature
o
f the order are the suctorial mouthparts, which are remarkabl
y
similar in all except a ver
y
f
ew mem

b
ers o
f
t
h
e group. T
h
e man
dibl
es an
d
max
ill
ae
f
orm two pa
i
rs o
f
p
i
erc
i
ng sty
l
ets
t
h
at are conta
i

ne
d
w
i
t
hi
nt
h
e
fl
ex
ibl
e, segmente
dl
a
bi
um (see C
h
apter 3, Sect
i
on 3.2.2 an
d
Fig
ure 3.17). T
h
est
yl
ets are somet
i
mes muc

hl
on
g
er t
h
an t
h
e
l
a
bi
um an
d
,w
h
en not
i
n use,
m
a
y
be coiled within an inte
g
umental fold, the crumena. Maxillar
y
and labial palps ar
e
211
THE HEMIPTER
O

ID
ORDER
S
absent, thou
g
h the labium, which never enters the tissue that is pierced, has a number of
sensor
y
hairs at its tip. Mouthparts are absent in male and man
y
female Coccoidea and th
e
sexual forms of some a
p
hids.
I
n Heteroptera t
h
e pronotum
i
s
l
arge, t
h
e mesonotum an
d
metanotum sma
ll
.Inmos
t

h
omopterans t
h
e pronotum
i
s sma
ll
an
d
co
ll
ar
lik
e, t
h
e mesonotum
i
swe
ll d
eve
l
ope
d
,t
he
metanotum somew
h
at
l
ess so. Usua

lly
two pa
i
rs o
f
w
i
n
g
s are present, t
h
ou
gh b
rac
hy
pter
y
and apter
y
are common, sometimes occurrin
g
in the same species. In homopterans the win
gs
are
g
enerall
y
held rooflike over the bod
y
; the fore win

g
s are of uniform texture and are often
o
f
a
h
ar
d
er cons
i
stency t
h
an t
h
e
hi
n
d
pa
i
r. In most Heteroptera t
h
ew
i
ngs are
h
e
ld fl
at, an
d

th
ere
i
s typ
i
ca
ll
y a mar
k
e
d diff
erence
i
nt
h
e cons
i
stency o
f
t
h
e
f
ore an
dhi
n
d
w
i
ngs. T

h
e
f
ore w
i
ngs (
h
eme
l
ytra) usua
ll
y are we
ll
sc
l
erot
i
ze
db
asa
ll
y, w
i
t
h
on
l
y a sma
ll di
sta

l
port
i
on
remainin
g
membranous. The hind pair remain membranous and, at rest, are folded beneat
h
t
he hemel
y
tra. The le
g
s are
g
enerall
y
identical and suited for a cursorial habit. However, in
certain groups they are modified for a variety of functions, such as catching prey, swimming
,
j
ump
i
ng, mov
i
ng over t
h
e water sur
f
ace, an

d
pro
d
uct
i
on o
f
soun
d
,ort
h
ey may
b
e vest
i
g
i
a
l
or a
b
sent
i
n
f
ema
l
es t
h
at are se

d
entary.
I
n its least specialized condition the abdomen has 11 se
g
ments of which the first two ma
y
b
e modified in connection with sound production, the ei
g
hth and ninth possess the external
genitalia, and the last two are extremely reduced. Frequently reduction of up to three anterior
segments occurs. T
h
eov
i
pos
i
tor
i
s comp
l
ete
i
n many
h
omopterans an
d
t
h

ose Heteroptera
th
at
l
ay eggs
i
np
l
ant t
i
ssues,
b
ut re
d
uce
d
or a
b
sent
i
not
h
er groups. T
h
e
h
omo
l
og
i

es o
f
t
he
male
g
enitalia are complex. Primitivel
y
, the enlar
g
ed ninth sternum bears a pair of lateral
claspers, while the cavit
y
formed b
y
inva
g
ination of the membrane between the ninth and
t
enth sterna contains the aedea
g
us and a pair of parameres. However, either the claspers or
b
ot
h
t
h
ec
l
aspers an

d
parameres are a
b
sent
i
n some groups.
T
h
ea
li
mentary cana
l
presents a w
id
evar
i
ety o
f
structura
l
mo
difi
cat
i
ons assoc
i
ate
d
w
i

t
h
th
e
li
qu
id di
et o
f
t
h
eor
d
er. Poster
i
or to t
h
ec
ib
ar
i
a
l
suc
ki
n
g
pump, w
hi
c

hi
s not part o
f
t
h
e
alimentar
y
canal, is a short fore
g
ut. The mid
g
ut is lar
g
e, frequentl
y
occup
y
in
g
ama
j
or part
of the abdomen and usuall
y
differentiated into several re
g
ions, of which the most anterior
i
s a crop

lik
e structure. T
h
e poster
i
or reg
i
on
i
stu
b
u
l
ar an
df
rom
i
t,
i
n many Heteroptera
,
ar
i
se many ceca t
h
at conta
i
n sym
bi
ot

i
c
b
acter
i
a. In Ap
hid
o
id
ea a spec
i
a
l
structure (myce-
t
ome) w
i
t
hi
nt
h
e
b
o
d
ycav
i
ty
h
ouses sym

bi
ot
i
c
b
acter
i
a
(
B
uc
h
ner
a
s
pp.) t
h
at prov
id
et
he
h
ost with the essential amino acids unavailable in the diet (see Chapter 16, Section 5.1.2)
.
I
n man
y
plantsuckin
g
Hemiptera the hind

g
ut is lon
g
and convoluted. The anterior part of
t
he hindgut and the Malpighian tubules may come to lie alongside the swollen midgut,
th
us prov
idi
nga“s
h
ortcut” to t
h
e
hi
n
d
gut
f
or t
h
e
l
arge vo
l
umes o
fli
qu
id f
eces (see F

i
g-
u
re 1
6
.
6
). In a few Heteroptera the midgut ends blindly or is entirely separated from th
e
h
ind
g
ut. Four Malpi
g
hian tubules are the rule, but rarel
y
there are onl
y
three or two. In
Aphidoidea there are no Malpi
g
hian tubules, and the function of excretion is taken ove
r
b
y the midgut. The central nervous system is highly specialized, and discrete abdominal
gang
li
a never occur. In most Hem
i
ptera t

h
ere
i
sas
i
ng
l
e compos
i
te ventra
l
gang
li
on;
in
ot
h
ers up to
f
our ventra
l
gang
li
a (su
b
esop
h
agea
l
, prot

h
orac
i
c, mesot
h
orac
i
c, an
d
compos
i
t
e
metat
h
orac
i
c-a
bd
om
i
na
l
)ma
yb
e
f
oun
d
. Usua

lly
eac
h
ovar
y
compr
i
ses
b
etween
f
our an
d
ei
g
ht acrotrophic ovarioles arran
g
ed in a
g
roup at the distal end of the lateral oviduct. In a few
h
omopterans, for example, aphids, there ma
y
, however, be up to 100 ovarioles. Paired acces-
sory g
l
an
d
san
d

zero to t
h
ree spermat
h
ecae are a
l
so present. Eac
h
test
i
s conta
i
ns
f
rom on
e
t
oseven
f
o
lli
c
l
es, w
hi
c
h
may or may not
b
e enc

l
ose
di
na
f
o
lli
cu
l
ar s
h
eat
h
.We
ll d
eve
l
ope
d
accessor
ygl
an
d
sa
l
so occur
i
nma
l
es. Var

i
ous t
y
pes o
f
su
b
cutaneous
gl
an
d
s are common
212
CHAPTER
8
i
n Hemiptera. Most Heteroptera possess scent
g
lands in both
j
uvenile and adult sta
g
es. In
j
uveniles these are t
y
picall
y
on the dorsal surface of abdominal se
g

ments 4–7. Thou
g
hin
some heteropterans the
g
lands of
j
uveniles continue to function in the adult sta
g
e, in most
spec
i
es new, ventra
ll
y pos
i
t
i
one
d
metat
h
orac
i
cg
l
an
d
s
diff

erent
i
ate at ec
l
os
i
on. Most
l
yt
h
e
g
l
an
d
s secrete repugnator
i
a
l
c
h
em
i
ca
l
s;
i
n some spec
i
es,

h
owever, aggregat
i
on, warn
i
ng, o
r
sexua
l
p
h
eromones are pro
d
uce
d
. Wax-secret
i
n
ggl
an
d
s occur
i
n man
y
Sternorr
hy
nc
h
aan

d
Ful
g
oroidea, usuall
y
on the dorsal surface of the abdomen. Their function is presumabl
y
p
rotective
.
Lif
eH
i
story and Hab
i
t
s
Homopterans and the ma
j
orit
y
of Heteroptera are terrestrial insects. Other Heteropter
a
show var
y
in
g
de
g
rees of adaptation to an aquatic existence. Some occur in the littoral or

i
ntertidal zone of the seashore, on marshy ground, or in damp moss. Others live on th
e
sur
f
ace o
f
water. Among t
h
e tru
l
y aquat
i
c
f
orms t
h
ere
i
s a range o
f
a
d
aptat
i
on
f
rom t
h
os

e
spec
i
es t
h
at must per
i
o
di
ca
ll
yv
i
s
i
tt
h
e sur
f
ace to resp
i
re (an
d
w
hi
c
h
per
i
o

di
ca
ll
y
fl
y
f
ro
m
o
ne location to another) to those that remain submer
g
ed permanentl
y
and respire b
y
means
o
fa
p
lastron (see Cha
p
ter 15, Section 4.2).
All homopterans and many Heteroptera feed on fluids from plants. All parts of the
pl
ant are attac
k
e
d
: roots, stem,

l
eaves,
fl
owers, an
d
see
d
s(o
f
ten w
h
en t
h
ese
h
ave
f
a
ll
en
to t
h
e groun
d
). Sternorr
h
ync
h
aan
d

Fu
l
goro
id
ea pr
i
nc
i
pa
ll
y
f
ee
di
np
hl
oem, C
i
ca
d
o
id
e
a
an
d
Cercopo
id
ea
i

nx
yl
em, an
d
Heteroptera
i
n parenc
hy
ma, w
hil
eC
i
ca
d
e
ll
o
id
ea var
yin
their choice of tissue
(
Carve
r
et al.
,
1991; Novotn
y
and Wilson, 1997). The remainin
g

H
eteroptera are predaceous, livin
g
on the bod
y
fluids of other arthropods and vertebrates.
It
i
spr
i
mar
il
y
b
ecause o
f
t
h
ese
f
ee
di
ng
h
a
bi
ts, ass
i
ste
di

n many cases
b
y extraor
di
nar
ily
hi
g
h
rates o
f
repro
d
uct
i
on (see Super
f
am
il
yAp
hid
o
id
ea), t
h
at t
h
eor
d
er

i
s cons
id
ere
db
y
m
an
y
peop
l
eto
b
et
h
e most econom
i
ca
lly i
mportant
i
nsect
g
roup. T
h
e
d
ama
gi
n

g
e
ff
ect on
plants ma
y
be direct or indirect. When insect populations are lar
g
e, the loss of sap result
s
i
n stunted
g
rowth and poor
y
ield and qualit
y
. Man
y
species, when feedin
g
,in
j
ect saliva that
c
auses necrosis of plant tissue. Indirectly, the effects are to weaken the plant, making it mor
e
suscept
ibl
e to attac

kb
yot
h
er pat
h
ogens, espec
i
a
ll
y
f
ung
i
an
d
v
i
ruses. Most
i
mportant o
f
a
ll,
h
owever,
i
st
h
ero
l

eo
f
Hem
i
ptera (espec
i
a
ll
yap
hid
s) as vectors o
f
v
i
ruses t
h
at cause ma
j
o
r
plant diseases, for example, mosaic, leaf roll,
y
ellows. Amon
g
the predaceous Hemiptera
,
af
ew
f
f

s
pecies ma
y
act as vectors for the transmission of disease; for example, certain Sout
h
A
merican Redu
v
iidae are carriers of Trypanosoma cruz
i
,a
fl
agellate protozoan that causes
trypanosom
i
as
i
s(C
h
agas’
di
sease), a
f
orm o
f
s
l
eep
i
ng s

i
c
k
ness among
h
umans. On t
he
b
ene
fi
c
i
a
l
s
id
e, many
h
omopterans p
l
ay an
i
mportant part
i
n wee
d
contro
l
;
f

or examp
l
e,
D
act
y
lopiu
s
species have been successfull
y
used in the control of the prickl
y
pear cactus
(
Op
unti
a
)
, and man
y
predaceous Heteroptera exert a ma
j
or controllin
g
effect on som
e
arthro
p
od
p

ests.
Th
ema
j
or
i
ty o
f
Hem
i
ptera are
bi
sexua
l
an
d
ov
i
parous. T
h
ere are,
h
owever, spec
i
e
s
t
h
at are
f

acu
l
tat
i
ve
l
y part
h
enogenet
i
c, an
d
some t
h
at are ovov
i
v
i
parous or v
i
v
i
parous. In
m
an
y
ap
hid
sa
ll

o
f
t
h
ese repro
d
uct
i
ve con
di
t
i
ons ma
yb
e met w
i
t
hi
nt
h
e same spec
i
es
i
n
the course of a
y
ear. In aphids partheno
g
enesis and viviparit

y
commonl
y
occur to
g
ether i
n
the sprin
g
and earl
y
summer, thus enablin
g
the insects to exploit full
y
the increased foo
d
a
v
a
il
a
bl
eatt
hi
st
i
me. Insem
i
nat

i
on
i
s typ
i
ca
ll
yo
f
t
h
e usua
li
ntragen
i
ta
l
type. However
,
i
nC
i
m
i
co
id
ea var
i
ous
f

orms o
fh
emocoe
li
c
i
nsem
i
nat
i
on occur (see C
h
apter 19, Sect
i
on
213
THE HEMIPTER
O
ID
ORDER
S
4
.3.1). The e
gg
s of homopterans are
g
enerall
y
simple, ovoid structures; those of Heteroptera
are ver

y
diverse in form and colorin
g
. The
y
are
g
lued to plant surfaces, inserted in crevice
s
or between ad
j
acent parts of a plant, laid in litter or soil, or in the case of predaceous species
,
i
nt
h
ev
i
c
i
n
i
ty o
f
a
h
ost
if
poss
ibl

e. In a
f
ew Heteroptera t
h
e eggs are stuc
k
on t
h
e
d
orsa
l
sur
f
ace o
f
ma
l
es. Parenta
l
care
i
ss
h
own
b
ya
f
ew spec
i

es. Usua
ll
y, t
h
ere are
b
etween t
h
ree
an
d
seven
i
nstars (a
l
most a
l
wa
y
ss
i
x
i
n Heteroptera). T
h
e
j
uven
il
e sta

g
es ten
d
to
f
ee
d
on t
h
e
same part of the plant as the adult, thou
g
h in some species
j
uveniles are found on the roots,
w
hile adults occur on the upper parts. Althou
g
h most Hemiptera are t
y
picall
y
exopter
yg
ot
e
i
nt
h
e

i
r postem
b
ryon
i
c
d
eve
l
opment, t
h
ere occurs
i
n some A
l
euro
did
ae an
d
w
i
nge
d
ma
l
e
Cocco
id
ea one or two rest
i

ng
i
nstars, t
h
e pupa
l
an
d
prepupa
l
stages. T
h
ese
i
nstars genera
ll
y
d
o not
f
ee
db
ut un
d
ergo some
d
egree o
f
metamorp
h

os
i
s.As
i
m
il
ar p
h
enomenon occurs
in
T
h
y
sanoptera
.
Phylogeny and Classification
L
ik
et
h
e Psocoptera, t
h
e Hem
i
ptera are a very anc
i
ent group w
h
ose
f

oss
il
recor
d
ex-
t
ends into the Lower Permian period. The earliest known Hemiptera are the Archesc
y
tinidae
(Fi
g
ure 2.9B) from the Lower Permian of Moravia (Czech Republic). These represent “stem-
group” homopterans, close to but not within the Auchenorrhyncha, which appear slightl
y
l
ater. Sternorr
h
ync
h
aan
d
Heteroptera
fi
rst appear
i
nt
h
e
f
oss

il
recor
d
o
f
t
h
e Upper Perm
i
an
.
B
yt
h
e Upper Perm
i
an, t
h
e auc
h
enorr
h
ync
h
ans were c
l
ear
l
y
di

v
i
s
ibl
e
i
nto Fu
l
goromorp
ha
and Cicadomorpha, and fossils assi
g
nable to extant families have been obtained from Upper
T
riassic and Lower Jurassic deposits. Among the Sternorrhyncha, the Psylloidea and their
T
T
sister
g
roup, Aleurodoidea, date from the Upper Permian, while the earliest Aphidoidea an
d
Cocco
id
ea occur
i
nt
h
eTr
i
ass

i
c. T
h
eCo
l
eorr
h
ync
h
a, w
h
ose
f
oss
il
recor
d
exten
d
s
b
ac
k
t
o
th
e Lower Jurass
i
c, s
h

are c
h
aracters w
i
t
hb
ot
h
t
h
e Auc
h
enorr
h
ync
h
aan
d
t
h
e Heteroptera
,
th
ou
gh
t
h
e consensus
i
st

h
at t
h
e
yb
ranc
h
e
d
o
ff
t
h
e
h
eteropteran
li
ne at an ear
ly d
ate, per
-
h
aps even in the Permian. Some authors consider this
g
roup sufficientl
y
distinct from othe
r
Hemi
p

tera as to warrant the rank of suborder. In the Jurassic the Hetero
p
tera underwent
a
wid
era
di
at
i
on, an
d
t
h
e comparat
i
ve
l
yr
i
c
hf
oss
il
recor
df
rom t
hi
s per
i
o

di
nc
l
u
d
es represen
-
t
at
i
ves o
f
many extant
f
am
ili
es, espec
i
a
ll
y Nepomorp
h
aan
d
Leptopo
d
omorp
h
a. A
l

t
h
oug
h
some
f
oss
il
C
i
m
i
comorp
h
a
h
ave
b
een o
b
ta
i
ne
df
rom t
h
e Upper Jurass
i
c, t
hi

s group an
d
its sister
g
roup, Pentatomorpha, underwent a ma
j
or expansion in the Cretaceous, paral
-
lelin
g
the radiation of the an
g
iosperms. A proposed ph
y
lo
g
en
y
of the order is shown i
n
F
igure 8.4
.
As note
di
nt
h
e
i
ntro

d
uct
i
on to t
hi
sor
d
er, t
h
e tra
di
t
i
ona
l
su
bdi
v
i
s
i
on o
f
t
h
e Hem
i
ptera
i
nto Homoptera an

d
Heteroptera
i
sno
l
onger tena
bl
e, t
h
e Sternorr
h
ync
h
anow
b
e
i
ng con
-
sidered the sister
g
roup to the rest of the Hemiptera (i.e., Auchenorrh
y
nch
a
+
H
eteroptera,
includin
g

Coleorrh
y
ncha) (see Carve
r
et al.
, 1991; Cam
p
bel
l
et al.
,
1994
;
S
orensen
et al.
,
1995; von Dohlen and Moran, 1995). Until recentl
y
, based on morpholo
g
ical criteria, it was
wid
e
l
y accepte
d
t
h
at t

h
et
h
ree su
b
or
d
ers were monop
h
y
l
et
i
c. Mo
l
ecu
l
ar stu
di
es
h
ave sup
-
porte
d
t
h
e monop
h
y

l
yo
f
t
h
e Sternorr
h
ync
h
aan
d
Heteroptera,
b
ut strong
l
y suggest t
h
at t
he
Auc
h
enorr
hy
nc
h
a
i
s a parap
hyl
et

i
c
g
roup. T
h
us, Sorense
n
et al.
(
199
5
) proposed the division
of Auchenorrh
y
ncha into two suborders, correspondin
g
to the Ful
g
oromorpha and Cicado
-
morpha of morpholo
gy
-based classifications. Within the Heteroptera, ei
g
ht infraorders ar
e
recogn
i
ze
di

nt
h
ec
l
ass
ifi
cat
i
on use
dh
ere (
l
arge
l
y
f
rom Carve
r
e
ta
l
., 1991). O
f
t
h
ese, t
he
G
erromorp
h

a correspon
d
to t
h
e Amp
hibi
cor
i
sae (amp
hibi
ous
b
ugs), t
h
e Nepomorp
h
ato
214
CHAPTER
8
F
IGURE 8.4.
A
p
ropose
d
p
h
y
l

ogeny o
f
t
h
e Hem
i
ptera. [A
f
ter M. Carver, G. F. Gross. an
d
T. E. Woo
d
war
d,
1
991, Hem
ip
tera,
i
n: T
h
e Insects of Austra
l
i
a
,2n
d
e
d
., Vo

l
. 1 (CSIRO, e
d
.), Me
lb
ourne Un
i
vers
i
t
y
Press. B
y
permission of the Division of Entomology, CSIRO.
]
t
h
eHy
d
rocor
i
sae (aquat
i
c
b
ugs), a
l
so
k
nown as Cryptocerata on t

h
e
b
as
i
so
f
t
h
e
i
rs
h
ort,
c
oncea
l
e
d
antennae, an
d
t
h
eC
i
m
i
comorp
h
a

+
Pentatomorp
h
atot
h
e Geocor
i
sae (terrestr
i
a
l
f
orms) of earlier s
y
stems.
215
THE HEMIPTER
O
ID
ORDER
S
F
I
GU
RE 8.5. Ps
yll
o
id
ea. T
h

e app
l
e suc
k
er,
P
sy
ll
ama
l
i
(
Psyllidae). [From A. D. Imms, 1957
,
A
G
eneral Textbook
of Entomo
l
og
y
,9t
h
e
d
. (rev
i
se
db
yO.W.R

i
c
h
ar
d
san
d
R. G
.
Dav
i
es
)
, Met
h
uen an
d
Co.]
S
uborder Sternorrhynch
a
L
ik
e Auc
h
enorr
hy
nc
h
a, Sternorr

hy
nc
h
a
h
aveanop
i
st
h
o
g
nat
h
ous
h
ea
d
w
i
t
h
out a
g
u
l
a
,
a small pronotum, and fore win
g
s (when present) with a uniform texture and held rooflike

o
v
er
the bod
y
at rest. Derived features of recent sternorrh
y
nchans include the position o
f
t
he base of the proboscis (between or posterior to the fore coxae), two-segmented tarsi
,
no mesot
h
orac
i
c troc
h
ant
i
n,asaw
lik
e egg-
b
urster, an
d
var
i
ous w
i

ng
f
eatures, nota
bl
yt
h
e
a
b
sence o
f
a vannus an
d
vanna
lf
o
ld i
nt
h
e
hi
n
d
w
i
ng. Anot
h
er
f
eature t

h
at
di
st
i
ngu
i
s
h
es
t
hem from auchenorrh
y
nchans is their multise
g
mented, filiform antennae
.
Super
f
ami
l
y Psy
ll
oi
d
ea
A
b
out 1400 o
f

t
h
e 2200 spec
i
es o
f
Psy
ll
o
id
ea are
i
nc
l
u
d
e
di
nt
h
e
f
am
il
y PSYLLIDAE.
T
he jumping plant lice, as members of the family are commonly called, are small (2–
5
m
m

l
on
g
)an
d
resem
bl
em
i
n
i
ature c
i
ca
d
as. T
h
e
yh
ave stron
ghi
n
dl
e
g
s, an
d
w
i
n

g
s are presen
t
in both sexes. The famil
y
contains man
y
pest species, which ma
y
cause
g
alls or a
g
enera
l
stuntin
g
of host plants. The commonl
y
held view that this stuntin
g
is caused b
y
viruses fo
r
whi
c
h
psy
llid

s serve as vectors
i
snow
b
e
li
eve
d
to
b
e wrong;
i
t
i
st
h
e tox
i
csa
li
va
i
n
j
ecte
d
d
ur
i
ng

f
ee
di
ng t
h
at causes t
h
e
d
amage. Examp
l
es o
f
pest spec
i
es are Ps
yll
ap
y
rico
la
an
d
P.
m
a
l
i (Fi
g
ure 8.

5
), two species introduced into North America from Europe, which feed o
n
pear and apple, respectivel
y
. TRIOZIDAE (650 species worldwide) have similar habits to
Ps
y
llidae, in which famil
y
the
y
are often included
.
Superfamily Aleurodoidea
I
nc
l
u
d
e
di
nt
h
eA
l
euro
d
o
id

ea
i
sas
i
ng
l
e
f
am
il
y, ALEURODIDAE (ALEYRODIDAE),
with approximatel
y
1100 species, commonl
y
known as whiteflies (Fi
g
ure 8.
6
). The
y
are
small insects, usuall
y
3 mm or less in len
g
th,
g
enerall
y

covered with a whitish wax
y
secre
-
t
ion. The
y
are commonl
y
found on the underside of leaves. The
g
roup is mainl
y
tropical o
r
su
b
trop
i
ca
l
,t
h
oug
h
a
f
ew spec
i
es are pests o

f
green
h
ouse crops
i
n temperate reg
i
ons. T
h
e
lif
ecyc
l
e
i
s comp
l
ex, an
d
part
h
enogenes
i
s
i
s common
l
y
i
nvo

l
ve
d
. Larvae are se
d
entary, an
d
th
e
fi
na
l
sta
g
e (“pupa”)
d
oes not
f
ee
db
ut un
d
er
g
oes a mar
k
e
d
metamorp
h

os
i
stot
h
ea
d
u
l
t.
Super
f
ami
l
y Coccoi
d
ea
S
ome 20 families are included in the Coccoidea, a large, heterogeneous group of more
th
an 7000 spec
i
es. Desp
i
te t
hi
s
h
eterogene
i
tyaun

if
y
i
ng c
h
aracter
i
st
i
co
f
t
h
e group
i
st
he
more or
l
ess
d
egenerate
f
ema
l
es. T
h
ese are apterous, an
d
may

b
e sca
l
e
lik
e, ga
ll
-
lik
e, or cov
-
ered with a wax
y
or powder
y
coatin
g
. For this reason the
y
are commonl
y
known as scale
2
1
6
CHAPTER
8
F
I
G

URE 8.6
.
Al
euro
d
o
id
ea. T
h
e
g
reen
h
ouse w
hi
te
fly
, Tria
l
euro
d
es vaporario-
r
um
(Aleurodidae). [From L. Llo
y
d, 1922, The control of the
g
reenhouse whitefl
y

(Asterochiton va
p
orariorum)
w
ith notes on its biology, Ann. A
pp
l. Biol
.
9
:
1
–
32
.
By
perm
i
ss
i
on o
f
t
h
e Assoc
i
at
i
on o
f
App

li
e
d
B
i
o
l
o
gi
sts.
]
i
nsects or mealybugs. Adult males are either apterous or have fore wings only, and have non
-
f
unct
i
ona
l
mout
h
parts. Fema
l
es are ov
i
parous (
i
nw
hi
c

h
case t
h
e eggs are usua
ll
y reta
i
ne
d
wi
t
hi
nt
h
e sca
l
y cover
i
ng o
f
a
f
ema
l
e), ovov
i
v
i
parous, or v
i

v
i
parous. Part
h
enogenes
i
s
is
c
ommon and hermaphroditism is known to occur in one
g
enus. Man
y
species have become
c
osmopolitan as a result of distribution b
y
trade. Notes on some of the commoner families
are
g
iven below. The DIASPIDIDAE (armored scales) form the lar
g
est famil
y
of coccoids
w
ith 2
5
00 species worldwide. Females are covered with a hard, waxy layer that is separate
f

rom t
h
e
b
o
d
y. Inc
l
u
d
e
di
nt
h
e
f
am
il
y are many pests o
f
trees an
d
s
h
ru
b
s,
f
or examp
l

e, Le
p
-
i
d
osa
ph
es u
l
m
i
,
t
h
eo
y
sters
h
e
ll
sca
l
e(F
ig
ure 8.7A,B), an
d
Q
ua
d
ras

p
i
d
iotus
p
erniciosu
s
,
the San Jose scale. Another lar
g
e famil
y
is the COCCIDAE (soft scales, wax scales) (1200
species), the female members of which show a wide ran
g
e of form. The famil
y
contains
severa
l
pests t
h
at are now w
id
esprea
d
t
h
roug
h

commerce;
f
or examp
l
e, Pu
lv
inaria innu
-
mera
b
i
l
i
s
,
t
h
e cottony map
l
e sca
l
e,
i
ntro
d
uce
df
rom Europe,
i
snow

f
oun
d
t
h
roug
h
out Nort
h
A
mer
i
ca
f
ee
di
n
g
on
f
orest, s
h
a
d
e, an
df
ru
i
t trees. T
h

e KERRIIDAE (LACCIFERIDAE),
a
small (70 species), mainl
y
tropical and subtropical
g
roup, have females that are extremel
y
de
g
enerate and live in a resinous cell.
L
acci
f
er lacc
a
,
the Indian lac insect,
p
roduces a secre-
tion from which shellac is
p
re
p
ared. The PSEUDOCOCCIDAE (2000 s
p
ecies worldwide)
are t
h
e common mea

l
y
b
ugs, so-ca
ll
e
db
ecause
f
ema
l
es are covere
d
w
i
t
h
a mea
l
yor
fil
amen-
tous waxy secret
i
on. Severa
l
spec
i
es o
f

Ps
eu
d
ococcu
s
(
F
i
gure 8.7C) are ma
j
or pests as t
h
e
y
are vectors of disease-causin
g
viruses. The ERIOCOCCIDAE (felt scales), a cosmopolitan
g
roup of some 500 species, were formerl
y
included in the Pseudococcidae because of their
c
lose resemblance to mealybugs. The family contains a number of pest species, includin
g
some ga
ll f
ormers, as we
ll
as some potent
i

a
ll
y
b
ene
fi
c
i
a
l
ones. Var
i
ou
s
D
act
yl
opius spec
i
es
,
f
or examp
l
e,
h
ave
b
een
i

ntro
d
uce
di
nto Austra
li
a
i
n an attempt to contro
l
t
h
epr
i
c
kl
y pear
c
actus
(
Op
untia) (see Chapter 24, Section 2.3). The MARGARODIDAE (
g
round pearls
)
(
250 species) form a widel
y
distributed famil
y

, included in which is
I
cerya purchasi
,
the
c
ottony-cushion scale (Figure 8.7D), an Australian species that was transplanted through
c
ommerce to many reg
i
ons o
f
t
h
ewor
ld
w
h
ere
i
t
b
ecame an
i
mportant pest o
f
c
i
trus
f

ru
i
t
.
In Ca
lif
orn
i
at
hi
s pest
h
as
b
een contro
ll
e
d
success
f
u
ll
y
f
o
ll
ow
i
ng t
h

e
i
ntro
d
uct
i
on o
f
t
h
e
pre
d
aceous
b
eet
le
Ro
d
o
l
ia car
d
ina
l
i
s
(see C
h
apter 24, Sect

i
on 2.3)
.
S
uperfamily Aphidoidea
M
ore t
h
an 4700 spec
i
es o
f
Ap
hid
o
id
ea
h
ave
b
een
d
escr
ib
e
d
,
i
nc
l

u
di
ng some o
f
t
h
e
w
orld’s most important insect pests. Some authors include all Aphidoidea in one famil
y
217
THE HEMIPTER
O
ID
ORDER
S
F
I
GU
RE 8.7
.
Coccoidea. (A) The oystershell scale
,
Le
p
idosa
p
hes ulmi (Diaspididae); (B) female
L. ul
m

i
;
(
C)
f
ema
l
e
l
ong-ta
il
e
d
mea
l
y
b
ug
,
P
seu
d
ococcus
l
ongispinu
s
(Pseu
d
ococc
id

ae); an
d
(D) ma
l
e cottony-cus
hi
on
s
ca
l
e
,
Icerya purc
h
asi
(
Mar
g
arodidae). [A, B, from D. J. Borror, D. M. Delon
g
, and C. A. Triplehorn, 197
6,
A
n
Introduction to the Study o
f
Insect
s
, 4th ed. By permission of Holt, Rinehart and Winston, Inc. C, D, from P P.
Grass´e (ed.), 1951

,
Tr
ait
r
r
ede
´
Z
oo
l
ogie,
V
ol. X. By permission of Masson, Paris.]
VV
whil
eot
h
ers recogn
i
ze up to 11
f
am
ili
es. Mem
b
ers o
f
t
h
e super

f
am
il
y are c
h
aracter
i
ze
db
y
th
e
i
r comp
l
ex
lif
ecyc
l
es,
i
nw
hi
c
h
t
h
e spec
i
es ta

k
es on a var
i
ety o
ff
orms an
df
requent
l
y
alternates between two taxonomicall
y
distinct host plants. B
y
far the lar
g
est of the fou
r
families of A
p
hidoidea, with some 4300 s
p
ecies, is the APHIDIDAE (a
p
hids,
p
lant lice,
greenfly, and black fly). Most aphids are found on leaves, shoots, or buds, though a few
spec
i

es
li
ve
i
n rat
h
er spec
i
a
li
ze
d
s
i
tuat
i
ons,
f
or examp
l
e,
i
nun
f
o
ld
e
dl
eaves or
i

n eart
h
s
h
e
l
ters espec
i
a
ll
y constructe
df
or t
h
em
b
y ants w
i
t
h
w
hi
c
h
t
h
ey are assoc
i
ate
d

. Most spec
i
es
are pol
y
morphic in different
g
enerations and reproduce in a variet
y
of wa
y
s. The
y
ma
y
also
show host alternation. A t
y
pical life c
y
cle, that of Dysaphis plantaginea, the ros
y
appl
e
aphid, is shown in Figure 8.8. Usually aphids overwinter in the egg stage. In spring the egg
s
h
atc
h
an

d
g
i
ve r
i
se to w
i
ng
l
ess, v
i
v
i
parous part
h
enogenet
i
c
f
ema
l
es. A var
i
a
bl
e num
b
er o
f
suc

h
generat
i
ons occur, an
d
t
h
ese are
f
o
ll
owe
db
yt
h
e pro
d
uct
i
on o
f
w
i
nge
di
n
di
v
id
ua

l
s
t
hat mi
g
rate to the alternate host on which reproduction continues. Later in the seaso
n
sexual males and females are produced, and the aphids return to the ori
g
inal host. The
y
mate and females la
y
the overwinterin
g
e
gg
s. Partheno
g
enesis provides the means b
y
which
ap
hid
s can
i
ncrease t
h
e
i

r popu
l
at
i
on extreme
l
y rap
idl
y. Fortunate
l
y, t
h
e occurrence o
f
a
l
arge
num
b
er o
f
pre
d
ators an
d
a
d
verse weat
h
er con

di
t
i
ons usua
ll
y
k
eep t
h
e
i
r num
b
ers
i
nc
h
ec
k.
Ap
hid
s
i
nsu
ffi
c
i
ent num
b
ers ma

yh
ave a
di
rect e
ff
ect on t
h
e
i
r
h
osts, caus
i
n
g
w
il
t
i
n
g
and stunted
g
rowth. The
y
are, however, economicall
y
more important throu
g
h their rol

e
218
CHAPTER
8
F
I
G
URE 8.8
.
L
if
ec
y
c
l
eo
f
t
h
e ros
y
app
l
eap
hid
,
D
ysap
h
is p

l
antaginea
.
[
A
f
ter D. J. Borror, D. M. De
l
on
g,
and C. A. Tri
p
lehorn, 1976,
A
n Introduction to the Study o
f
Insects
,
4th ed. B
y
permission of Holt, Rinehart and
Winston, Inc.
]
as vectors o
fdi
sease-pro
d
uc
i
ng v

i
ruses. M
i
gratory spec
i
es t
h
at are not part
i
cu
l
ar
l
y
h
ost
-
spec
ifi
c are espec
i
a
ll
y
i
mportant, s
i
nce t
h
ese can transm

i
t
di
seases among a w
id
evar
i
ety o
f
p
l
ants. M
y
zus persica
e
,
t
h
e
g
reen peac
h
ap
hid
(F
ig
ure 8.9A),
i
st
h

ec
l
ass
i
c examp
l
e,
b
e
i
n
g
k
nown as a vector for more than 100 virus diseases
.
P
EMPHIGIDAE (ERIOSOMATIDAE) are closel
y
related to the Aphididae. The famil
y
i
s small (275 species), widely distributed (often by commerce), and includes both above- and
b
e
l
ow-groun
df
ee
d
ers an

d
many ga
ll
-ma
ki
ng spec
i
es. Many o
f
t
h
ewoo
ll
yap
hid
s, so-ca
ll
e
d
g
,
yp , ]
219
THE HEMIPTER
O
ID
ORDERS
b
ecause of the woollike wax
y

filaments that the
y
produce, are included in this famil
y
. Mos
t
species under
g
o host alternation. The ADELGIDAE (CHERMIDAE) are a small famil
y
(50
species) whose members are confined to conifers. The
y
feed on needles, twi
g
s, or within
ga
ll
s. Most spec
i
es a
l
ternate
h
osts, t
h
epr
i
mary one a
l

ways
b
e
i
ng spruce. T
h
e PHYLLOX
-
E
RIDAE (70 spec
i
es) are a sma
ll b
ut w
id
e
l
y
di
str
ib
ute
df
am
il
yt
h
at
i
nc

l
u
d
es a num
b
er o
f
pests,
f
or examp
l
e
,
V
iteu
s
(
=
Phyll
oxera
)
v
itifo
l
i
i
,
th
ev
i

ne or
g
rape p
hyll
oxera t
h
at
d
ev
-
astated European vine
y
ards in the 1870s and 1880s followin
g
its accidental introduction
on North American vines. In North America win
g
ed females o
f
V
.
v
iti
f
oli
i
p
roduce two
t
ypes o

f
o
ff
spr
i
ng, ga
lli
co
l
ae t
h
at
f
orm
l
ea
f
ga
ll
s(F
i
gure 8.9B,C) an
d
ra
di
c
i
co
l
ae t

h
at
li
v
e
u
n
d
ergroun
df
ee
di
ng on t
h
ev
i
ne roots. In Europe on
l
yra
di
c
i
co
l
ae are pro
d
uce
d
. Bot
h

type
s
o
f
o
ff
spr
i
ng repro
d
uce part
h
enogenet
i
ca
ll
yt
h
roug
h
out t
h
e summer. Roots o
f
t
h
e natura
l
N
orth American hosts are resistant to attack b

y
V
. vitifolii
w
hereas those of the Europea
n
vine are hi
g
hl
y
susceptible. Successful control of the pest in Europe was thus achieved b
y
growing the European vines on resistant North American rootstock
.
S
u
b
or
d
er Auc
h
enorr
hy
nc
h
a
The two features that characterize this suborder are the complex tympanal organs an
d
antennae w
h

ose
fl
age
ll
um
i
sar
i
sto
id
(
h
a
i
r
lik
e). T
h
e
l
a
bi
um or
i
g
i
nates
f
rom t
h

e poster
i
or
h
ea
d
reg
i
on, an
d
t
h
ere
i
snogu
l
a. Except
f
or t
h
ose o
f
C
i
ca
d
o
id
ea, t
h

e
hi
n
dl
egs are sp
i
ne
d
and modified for
j
umpin
g.
I
n
f
raorder Fulgoromorph
a
Superfamily Fulgoroide
a
I
nt
h
e
l
ar
g
e,
h
etero
g

eneous super
f
am
ily
Fu
lg
oro
id
ea, w
h
ose mem
b
ers are common
ly
known as plant hoppers, there are more than 10,000 species arran
g
ed in some 20 families.
Plant hoppers are mainl
y
phloem feeders on hi
g
her plants, but a few feed on fun
g
i. Th
e
majority of species are 1 cm or less in length, though some tropical forms may reach a length
of
5
cm or more. Some of the larger and commonly encountered families are as follows.
Th

e CIXIIDAE (F
i
gure 8.10A), w
i
t
h
more t
h
an 1300 spec
i
es
di
str
ib
ute
d
t
h
roug
h
out t
h
e
w
orld, are re
g
arded as the most primitive ful
g
oroid famil
y

. Little is known of their biolo
gy,
but
the
y
oun
g
sta
g
es are subterranean and t
y
picall
y
feed on
g
rass roots. Larvae of a few
s
p
ecies have been encountered in ant nests while others are cave dwellers with reduce
d
eyes. DELPHACIDAE (F
i
gure 8.10B) are sma
ll
(
l
ess t
h
an1cm
i

n
l
engt
h
)
f
u
l
goro
id
st
h
at
f
requent
l
y
h
ave re
d
uce
d
w
i
ngs. T
hi
s
i
st
h

e
l
argest
f
am
il
yo
ff
u
l
goro
id
s, an
di
ts more t
h
a
n
1
5
00 species are reco
g
nized b
y
the lar
g
e spur on the tibia of the hind le
g
s. A few members
of this famil

y
are serious pests b
y
actin
g
as vectors of virus diseases. The DERBIDAE
constitute a mainl
y
tropical famil
y
, whose more than 800 species t
y
picall
y
have ver
y
lon
g
wi
ngs an
df
ee
d
on
f
ung
i
or
hi
g

h
er p
l
ants. DICTYOPHARIDAE (F
i
gure 8.10C) are me
di
um
-
s
i
ze
d
Fu
l
goro
id
ea w
h
ose
h
ea
db
ears a
di
st
i
nct anter
i
or process. T

h
e
f
am
il
y, w
hi
c
h
conta
i
ns
more than
6
00 species, is widel
y
distributed throu
g
h arid or semiarid areas of the world. The
F
ULGORIDAE, with about 700 species, are a widel
y
distributed
g
roup, man
y
of whos
e
members are known as lantern flies because the inflated anterior
p

art of the head was
b
e
li
eve
d
or
i
g
i
na
ll
yto
b
e
l
um
i
nous. T
h
e
l
argest p
l
ant
h
oppers are mem
b
ers o
f

t
hi
s
f
am
il
y.
Th
e ACHILIDAE
f
ormaw
id
e
l
y
di
str
ib
ute
d
,t
h
oug
h
pr
i
mar
il
y trop
i

ca
lf
am
il
y conta
i
n
i
ng
a
b
out 380 spec
i
es. Juven
il
es
li
ve
b
eneat
hb
ar
k
or
i
n rott
i
n
g
woo

d
. More t
h
an 1000 spec
i
es
220
CHAPTER
8
F
I
GU
RE 8.10.
F
ul
g
oroidea. (A)
C
ixius an
g
ustatu
s
(
Cixiidae
)
;
(
B
)
S

tenocranus dorsalis
(
Del
p
hacidae); (C)
S
colo
p
s
p
erdix (Dictyopharidae); (D)
A
normenis se
p
tentrionali
s
(
Flatidae); and (E) Fitchiella robert
s
on
i
(Issidae).
[From H. Os
b
orn, 1938, T
h
eFu
lg
or
id

ae o
f
O
hi
o
,
O
h
io Bio
l
. Surv. Bu
ll.
6
(
6):283–349 (Bulletin #3
5
). B
y
permission
o
f the Ohio Biolo
g
ical Surve
y
.
]
are
i
nc
l

u
d
e
di
nt
h
e
f
am
il
y FLATIDAE (F
i
gure 8.10D), a
hi
g
hl
y spec
i
a
li
ze
d
,ma
i
n
l
y trop
i
ca
l

,
g
roup. Mem
b
ers o
f
t
hi
s
f
am
ily
resem
bl
e mot
h
s
by
v
i
rtue o
f
t
h
e
i
rtr
i
an
g

u
l
ar, opaque te
g
m
i
na,
w
hich are folded to form a steep roof over the bod
y
. The ISSIDAE (Fi
g
ure 8.10E) ar
e
another lar
g
e famil
y
, with more than 1100 species. Its members are mostl
y
dull colored and
f
requent
l
y
h
ave a squat,
b
eet
l

e
lik
e
f
ac
i
es.
In
f
raorder Cicadomorpha
S
uperfamily Cercopoidea
Th
e mem
b
ers o
f
t
hi
s sma
ll
, rat
h
er
h
omo
g
eneous super
f
am

ily
are arran
g
e
di
nt
h
ree
f
amilies, CERCOPIDAE, APHROPHORIDAE, and MACHAEROTIDAE. S
p
ecies are sel-
dom more than 15 mm in len
g
th and frequentl
y
strikin
g
l
y
colored as adults. The larvae
of
a
f
ew Cercop
id
ae are su
b
terranean,
b

ut most
l
yt
h
ey
li
ve e
i
t
h
er
i
n a mass o
ff
rot
h
(Cer-
c
op
id
ae an
d
Ap
h
rop
h
or
id
ae), w
h

en t
h
ey are
k
nown as cuc
k
oo-sp
i
t
i
nsects or sp
i
tt
l
e
b
ugs,
o
r
i
naca
l
careous tu
b
e
(
Mac
h
aerot
id

ae
)
.T
h
e
f
unct
i
on o
f
t
h
ese structures, w
hi
c
h
are a
f-
fixed to plant stems, is to provide protection from predators and to prevent desiccation
.
221
THE HEMIPTER
O
ID
ORDER
S
F
I
GU
RE 8.11.

C
ercopoidea. A fro
g
hopper
,
P
hilaenus s
p
umariu
s
(
Cer-
copidae). [From D. J. Borror, D. M. Delong, and C. A. Triplehorn, 1976
.
An
Intro
d
uction to t
h
e Stu
d
y of Insects
,
4t
h
e
d
.B
y
perm

i
ss
i
on o
f
Broo
k
s/Co
l
e,
ad
ivision of Thomson Learnin
g
.
]
A
d
u
l
ts are act
i
ve,
h
opp
i
ng
i
nsects t
h
at

i
n many cases
b
ear a cru
d
e resem
bl
ance to a
f
rog
,
h
ence t
h
e common name o
ff
rog
h
opper. Ap
h
rop
h
or
id
ae, w
i
t
h
1300 spec
i

es,
f
orm t
h
e
l
arges
t
f
am
ily
w
i
t
h
aw
id
e
di
str
ib
ut
i
on. Its mem
b
ers are usua
lly f
oun
d
on

h
er
b
aceous p
l
ants,
b
ut
af
ew
f
f
live on trees to which the
y
sometimes do considerable dama
g
e. Cercopidae (100
0
species) are also widespread but particularl
y
common in the tropics. A few species are
pests o
f
grasses an
d
c
l
overs,
f
or examp

l
e, P
h
i
l
aenus spumariu
s
,
t
h
e mea
d
ow sp
i
tt
l
e
b
ug
(F
i
gure 8.11). Mac
h
aerot
id
ae (100 spec
i
es) are restr
i
cte

d
to As
i
a, trop
i
ca
l
A
f
r
i
ca, an
d
Austra
li
a
.
Super
f
ami
l
y Cica
d
oi
d
e
a
C
icadas are common insects in all of the warmer re
g

ions of the world. The
y
are
g
enerall
y
b
etween 2 and
5
cm in length and are particularly well known because of their sound-
pro
d
uc
i
ng a
bili
t
i
es an
d
t
h
e
l
engt
h
o
f
t
i

me requ
i
re
df
or
j
uven
il
e
d
eve
l
opment. V
i
rtua
ll
ya
ll
o
f
t
he 1
5
00 or so species are placed in the famil
y
CICADIDAE. The sound-producin
g
or
g
an

s
(t
y
mbals) are located on the dorsal side of the first abdominal se
g
ment of males onl
y
. The
a
uditor
y
t
y
mpana are better developed in males than females and are found on the ventral
s
id
eo
f
t
h
e anter
i
or a
bd
om
i
na
l
segments. Most c
i

ca
d
as requ
i
re severa
l
years
f
or
j
uven
il
e
d
eve
l
opment. A we
ll
-
k
nown per
i
o
di
cc
i
ca
d
a
i

s Ma
g
icica
d
a septen
d
eci
m
(
F
i
gure 8.12)
,
whi
c
h
,
i
nt
h
e eastern Un
i
te
d
States, requ
i
res 17
y
ears
f

or
i
ts
d
eve
l
opment;
i
n contrast, t
h
e
southern form takes onl
y
13
y
ears to mature. A larva spends this entire period under
g
round,
feedin
g
on roots, especiall
y
those of perennial plants. Prior to the final molt the larva leaves
t
he soil to complete its metamorphosis on a tree or other object. Eggs are laid in twigs, a
process t
h
at may cause cons
id
era

bl
e
di
e
b
ac
k
o
f
t
h
e tree.
F
IGURE 8.12. C
i
ca
d
o
id
ea. Juven
il
e per
i
o
di
cc
i
ca
d
a

.
M
agi-
cica
d
a septen
d
ecim
(
C
i
ca
did
ae
)
. [From C. L. Mar
l
att, 1907,
The periodical cicada
,
Bull. U.S. Dept A
g
r. Bur. Ent. (N. S.
)
71
:181
pp.]
222
CHAPTER
8

F
IGURE 8.13. Cicadelloidea. (A) The beet leafhopper, Circuli
f
er tenellu
s
(
Cicadellidae); (B) the potato leafhop-
per,
Empoasca fa
b
ae (C
i
ca
d
e
llid
ae); an
d
(C) t
h
e
b
u
ff
a
l
o tree
h
o
pp

er,
S
tictocep
h
a
l
a
b
u
b
a
l
u
s
(
Mem
b
rac
id
ae
)
. [A, B,
f
rom D. M. Delon
g
, 1948, The leafhoppers, or Cicadellidae, of Illinois,
B
ull. Ill. Nat. Hist.
S
ur

v
.
2
4(2):97–376. B
y
permission of the Illinois Natural History Survey. C, from D. J. Borror, D. M. Delong, and C. A. Triplehorn, 1976
,
An Intro
d
uction to t
h
e Stu
d
y of Insects
,
4t
h
e
d
.B
y
perm
i
ss
i
on o
f
Broo
k
s/Co

l
e, a
di
v
i
s
i
on o
f
T
h
omson Learn
i
n
g
.
]
S
uperfami
l
y Cica
d
e
ll
oi
d
ea
M
ost o
f

t
h
eC
i
ca
d
e
ll
o
id
ea are p
l
ace
di
n two very
l
arge
f
am
ili
es, CICADELLIDAE (JAS-
S
IDAE) (
l
ea
fh
oppers) an
d
MEMBRACIDAE (tree
h

oppers). T
h
e cosmopo
li
tan C
i
ca
d
e
llid
a
e
(
20,000 species), which form the lar
g
est homopteran famil
y
, are found on almost all t
y
pe
s
o
f plants. The
y
rank second onl
y
to the Aphididae in the enormit
y
of their numbers and,
i

n consequence, are major pests. They cause a wide variety of injuries to plants. They ma
y
remove
l
arge quant
i
t
i
es o
f
sap,
bl
oc
k
t
h
ep
hl
oem tu
b
es, or
d
estroy t
h
ec
hl
orop
h
y
ll

so t
h
a
t
growt
hi
s stunte
d
. Many are vectors o
f
v
i
ruses t
h
at cause
di
sease. A
f
ew
d
amage t
h
ep
l
ants
b
y
their oviposition habits. Two well-known pests are
C
irculifer tenellus

,
the beet leafhopper
(
Fi
g
ure 8.13A), and
E
mpoasca faba
e
, the potato leafhopper (Fi
g
ure 8.13B), which feed
s
o
n solanaceous plants, beans, celer
y
, alfalfa, and various flowers. Several other species
of
Empoasc
a
are a
l
so
i
m
p
ortant
p
ests (see Swan an
d

Pa
pp
, 1972). Mem
b
rac
id
ae (2400
spec
i
es) are eas
il
y recogn
i
ze
db
yt
h
e
i
r enormous pronotum t
h
at pro
j
ects
b
ac
k
war
d
over t

h
e
a
bd
omen an
d
o
f
ten assumes
bi
zarre s
h
apes. T
h
e
f
am
ily i
spr
i
mar
ily
neotrop
i
ca
l
,T
h
e
y

are
g
enerall
yg
re
g
arious and attended b
y
ants for the hone
y
dew the
y
produce. The
y
are seldo
m
o
f economic im
p
ortance; however,
S
tictocephala bubalu
s
,
the buffalo treehopper (Fi
g
ur
e
8
.13C), may

d
amage young
f
ru
i
t trees an
d
nursery stoc
k
as a resu
l
to
fi
ts egg-
l
ay
i
ng act
i
v
i
ty.
Suborder Heteroptera
In Heteroptera the head is usuall
y
pro
g
nathous, almost alwa
y
s with a

g
ula; the pronotum
i
s well developed; and fore wings when present are in the form of hemelytra, with wings
h
e
ld fl
at over t
h
e
b
o
d
yw
h
en at rest
.
In
f
raorder Coleorrhyncha
S
uperfamily Peloridioidea
Contained in this superfamil
y
is a sin
g
le, Southern Hemisphere famil
y
, PELORIDI
-

IDAE, whose 25 species are small, flattened, cr
y
pticall
y
colored Hemiptera found amon
g
223
THE HEMIPTER
O
ID
ORDER
S
moss and liverworts or in caves, The famil
y
is all that remains of a formerl
y
numerous and
w
idespread Northern Hemisphere
g
roup, with fossils from as earl
y
as the Lower Jurassic pe-
riod, Their current distribution is dis
j
unct, with species occurrin
g
in South America and the
Austra
li

an reg
i
on. Pe
l
or
idiid
s
h
ave a
f
asc
i
nat
i
ng m
i
xture o
fh
omopteran an
dh
eteropteran
f
eatures, as we
ll
as suc
h
ancestra
l
c
h

aracters as t
h
e comp
l
ete tentor
i
um,
di
screte pro- an
d
mesot
h
orac
i
c
g
an
gli
a, an
d
e
igh
tpa
i
rs o
f
a
bd
om
i

na
l
sp
i
rac
l
es
.
Infraor
d
er Enicocep
h
a
l
omorp
h
a
Super
f
ami
l
y Enicocep
h
a
l
oi
d
ea
The 260 or so species are usually arranged in a single family, ENICOCEPHALIDAE,
whi

c
hh
as a pre
d
om
i
nant
l
y trop
i
ca
l
to su
b
trop
i
ca
ldi
str
ib
ut
i
on. T
h
ey are sma
ll
-tome
di
um-
s

i
ze
db
ugs, typ
i
ca
ll
y
f
oun
d
among
l
ea
fli
tter, rott
i
ng
l
ogs, etc., w
h
ere t
h
eypreyonot
h
e
r
arthropods. Man
y
are brach

y
pterous or apterous, thou
g
h win
g
ed species often appear i
n
lar
g
e swarms at dusk, hence their common name of
g
nat bu
g
s.
Infraorder Dipsocoromorpha
Superfamily Dipsocoroidea
About three quarters of the approximately 22
5
species of Dipsocoroidea are included i
n
th
e cosmopo
li
tan
f
am
ily
SCHIZOPTERIDAE. T
h
ese are m

i
nute, pre
d
aceous
i
nsects
f
oun
d
in dam
p
habitats such as moss, leaf litter, and ant nests.
Infraorder Gerromorpha
S
ome aut
h
ors
i
nc
l
u
d
ea
ll
mem
b
ers o
f
t
h

e
i
n
f
raor
d
er
i
nas
i
n
gl
e super
f
am
ily
, Gerro
id
ea
.
Others, includin
g
Carve
r
et al.
(1991), split the
g
roup into the followin
g
four superfamilies

.
Superfamily Mesovelioide
a
T
h
e 30 or so spec
i
es are arrange
di
nas
i
ng
l
e, w
id
e
l
y
di
str
ib
ute
df
am
il
y, MESOVELI-
I
DAE. These lon
g
-le

gg
ed, predaceous bu
g
s are usuall
y
found at the mar
g
ins of ponds,
crawlin
g
amon
g
ve
g
etation and debris; sometimes the
y
move over the water surface. A few
species occur away from water in damp places, including caves.
Super
f
amily Hebroidea
H
ebroidea are ver
y
small, semiaquatic bu
g
s found amon
g
moss or on ponds with a
n

abundance of floatin
g
or emer
g
ent ve
g
etation where the
y
pre
y
on small arthropods. The
group has a cosmopolitan distribution and its 150 species are included in a single family,
HEBRID
A
E.
Super
f
ami
l
y Gerroi
d
ea
A
ll b
ut a
b
out a
d
ozen o
f

t
h
e approx
i
mate
l
y 1000 spec
i
es
i
nt
hi
s super
f
am
il
y are
i
nc
l
u
d
e
d
i
n two equa
ll
ys
i
ze

df
am
ili
es, GERRIDAE an
d
VELIIDAE. Gerr
id
ae, common
l
y
k
nown as
pon
d
s
k
aters an
d
water str
id
ers (F
ig
ure 8.14A), are e
l
on
g
ate
b
u
g

sw
h
ose
b
o
dy i
s covere
d
w
ith a dense la
y
er of waterproof hairs. Members of this widespread
g
roup are mainl
y
foun
d